ADRIANO AFONSO SPIELMANN
Estudos taxonômicos em Parmotrema s.l.
{Par meliaceae, Ascomy cota liquenizados) com ácido salazínico
Tese apresentada ao Instituto de Botânica da
Secretaria do Meio Ambiente, como parte dos
requisitos exigidos para a obtenção do título de
DOUTOR em BIODIVERSIDADE VEGETAL E
MEIO AMBIENTE, na Área de Concentração de
Plantas A vasculares e Fungos.
SAO PAULO
2009
ADRIANO AFONSO SPIELMANN
Estudos taxonômicos em Parmotrema s.l.
{Par meliaceae, Ascomy cota liquenizados) com ácido salazínico
Tese apresentada ao Instituto de Botânica da
Secretaria do Meio Ambiente, como parte dos
requisitos exigidos para a obtenção do título de
DOUTOR em BIODIVERSIDADE VEGETAL E
MEIO AMBIENTE, na Área de Concentração de
Plantas A vasculares e Fungos.
ORIENTADOR: DR. MARCELO PINTO MARCELLI
Ficha Catalográfica elaborada pela Seção de Biblioteca do Instituto de Botânica
Spielmann, Adriano Afonso
S755e Estudos taxonômicos em Parmotrema s.l. (Parmeliaceae, Ascomycota
liquenizados) com ácido salazínico / Adriano Afonso Spielmann ~ São Paulo,
2009.
152p.il.
Tese (Doutorado) ~ Instituto de Botânica da Secretaria de Estado do Meio
Ambiente, 2009
Bibliografia.
1. Liquens. 2. Canomaculina. 3. Rimelia. I. Título
CDU: 582.29
"Gostar de algo é melhor do que
meramente conhecê-lo,
e encontrar alegria nesse algo
é melhor do que meramente gostar dele".
Confúcio, Os Analectos, Livro VI-20
AGRADECIMENTOS
A CAPES pela bolsa concedida no início do doutorado, e especialmente à FAPESP
(Processo n° 05/53740-5) pela bolsa concedida em seguida. Sem tais bolsas este trabalho não
teria sido possível.
Ao Instituto de Botânica de São Paulo (IBt), por possibilitar a vinda e permanência em São
Paulo por estes últimos seis anos, cinco dos quais se passaram enquanto morava no Alojamento
do Instituto.
Ao Programa do curso de Pós-graduação em Biodiversidade Vegetal e Meio Ambiente, do
IBt, especialmente à Dra. Solange C. Mazzoni- Viveiros, Dra. Sónia Machado de Campos
Dietrich e Sra. Márcia Regina Angelo (Marcinha). Obrigado pela oportunidade de ter sido seu
aluno "01.01.01" durante o mestrado e "01.02.01" durante o doutorado! É muito bom ver o
quanto o programa cresceu nos últimos anos, e faço votos para que tenha um futuro cada vez
melhor.
Ao meu orientador, Dr. Marcelo Pinto Marcelli, agradeço profundamente a oportunidade
de ter vivido seis anos intensos e produtivos, "respirando liquens" e aprendendo muito, o tempo
inteiro, com as coisas que "dão certo" e com aquelas que "dão errado" também. Obrigado por
todo o incentivo, cobrança e zelo, e por propiciar as todas as vivências que tivemos durante o
mestrado/doutorado, que sem dúvida são parte integral de nossa formação. "Lutar pelos liquens"
é um sonho que já se tornou realidade. Posso dizer que sairei não um liquenólogo perfeito, mas
um lutador vibrante! Obrigado também à Dra. Agnes Elisete Luchi, pela companhia sempre
serena e alegre.
A American Bryological and Lichenological Society - ABLS e International Association
for Lichenology - IAL, pelo custeio de algumas despesas para participar do IAL - 6 em
Asilomar, Califórnia, E.U.A. A FAPESP, por ter financiado o restante da viagem. E às pessoas
incríveis que tornaram possível esta participação em um congresso internacional, especialmente
o Dr. Thomas Nash III (Tom).
Aos curadores dos seguintes herbários, que gentilmente nos enviaram material para estudo:
(acrónimos conforme o Index Herbariorum): ASU, B, BAFC, BM, C, CANB, CHR, COLO,
CTES, DUKE, F, FI, G, H, K, L, LD. M, MIN, MSC, O, P, RO, S, TNS, TUR, UPS, UPCB, US,
W, e WU.
O herbário FH nos deu uma grande oportunidade de efetuar estudos em suas importantes
coleções. Agradeço à Dra. Emily Wood e à bibliotecária Judith A. Warnement e todo o seu
eficiente time, bem como ao Dr. Donald H. Pfister. Um agradecimento especial é devido à Dra.
Michaela Schmull, por todo o carinho, paciência e interesse em nossa pesquisa, bem como envio
e re-envio de fotos de importante material.
Estimados Robert Líicking & Eimy Rivas Plata, muchas gradas por todo o apoio e
dedicação que tornaram possível nossa estadia em Chicago (F)! Vocês são pessoas realmente
adoráveis! Aproveito também para agradecer ao Dr. Thorsten Lubsch por nos auxiliar nesta
visita.
Um agradecimento especial à Dra. Mónica Adler, Dra. Lidia Ferraro e Dra. Sionara
Eliasaro, pelo sempre rápido atendimento aos nossos pedidos.
Aos muitos liquenólogos que me enviaram bibliografia por todos estes anos, um
agradecimento muito sincero! Não posso deixar de mencionar os Drs. Robert Líicking, Teuvo
Ahti, Hiroyuki Kashiwadani, Philippe Clerc, John Elix, Klaus Kalb e Mónica Adler.
A bibliotecária do IBt, Maria Helena Simões Costa Fernandes Gallo, sempre manteve
portas abertas aos meus pedidos, auxiliando na procura e obtenção de muita, repito, muita
bibliografia antiga e importante. Não há jeito de deixar um "ratinho" mais feliz. Muito obrigado!
Em toda esta jornada foi um privilégio poder contactar e principalmente conhecer a grande
família liquenológica mundial. Graças ao GLAL pudemos conhecer nossos hermanos latino-
americanos, assim como liquenólogos de habitats mais distantes, a quem agradeço
profundamente toda simpatia e estímulo.
A Dra. Neli Kika Honda, muito obrigado pelo excelente curso que nos proporcionou em
Campo Grande. Espero ainda trabalharmos muito em conjunto.
Ao Dr. John Elix, muito por disponibilizar informações químicas e por se dispor a efetuar
análises de praticamente todos os tipos das espécies aqui tratadas.
A toda a Seção de Micologia e Liquenologia do IBt, muito obrigado por todo o apoio e
fornecimento de facilidades para a realização do trabalho. Em especial à Dra. Adriana Gugliotta,
pelo empréstimo da máquina fotográfica para que eu pudesse obter muitas das figuras de detalhe
que aparecem neste trabalho.
Ao Dr. Carlos Ernesto Schaeffer, da Universidade Federal de Viçosa (UFV), muitíssimo
obrigado por disponibilizar uma "enxada de trabalho" (estereomicroscópio), que auxiliou
muitíssimo durante todo o trabalho. Obrigado também por ceder o tempo livre na Antártica
(geralmente fortes nevascas) para que eu pudesse trabalhar na tese.
Ao colega Michel Benatti, também subcurador do herbário de liquens, muitíssimo
obrigado por toda a competência com que conduziu nossos empréstimos e devoluções de
material de herbário, buscando soluções a todos os problemas envolvidos (que não foram
poucos) e por fim sempre fazendo que o material chegasse em nossas mãos, são e salvo.
A querida Juçara Bordin (Jusci Musci), por todo o auxílio na triagem de espécimes, pelo
apoio e simpatia constantes, assim como à Priscila Silva (Pri) e à Regina Hirai (Rê), pela
presença sempre bem vinda. Também agradeço às meninas do "apartamento Lili", Berta
Villagra, Patrícia Junbluth e Luciana Canêz, por este convívio tão legal.
Aos amigos e amigas do "Orquídeas 24", Carlos Wetzel (Cati), Kleber dos Santos
(Klebér), Fernanda Ferrari (Fê) e Angélica Righetti (Angeliquinha), obrigado pelos bons
momentos que sempre passamos juntos.
Ao amigo do peito Luciano Bornholdt, pelo estímulo constante e leitura de algumas partes
do texto. Aos amigos Marcelo Aloísio Sulzbacher, Vagner Gularte Cortez e Gilberto Coelho,
pela força, bibliografia e amizade. Ao colega Marcos Junji Kitaura ("Marcos James"), pelo
auxílio nas coletas, discussões liqueno lógicas e companhia constante em congressos e encontros.
A colega Patrícia Jungbluth, muito obrigado pelo convite para várias coletas, pela presença
constante, e pelas discussões científicas que amiúde ocorrem. Obrigado também pelas leituras de
partes da tese e por ajudar na triagem de material e cromatografia.
A colega, amiga, companheira e noiva Luciana Canêz, uma pessoa especial que sempre
ajudou, o tempo inteiro, na leitura, correções, triagem de material, finalização da tese etc. Sou
duplamente feliz, por viver com liquens e contigo. Para mim, esta é a simbiose perfeita.
Dedico este trabalho aos meus pais, Seu Romeu e Dona íris, por
terem sempre acreditado em seu filho.
Adriano Afonso Spielmann
Sumário
Resumo
Abstract
Introdução 01
Histórico de Parmotrema s.l. com ácido salazínico 01
Parmotrema Massalongo 01
Canomaciãina Elix & Hale e Rimeliella Kurok 02
Rimelia Hale & Fletcher 02
Histórico das Espécies de Parmotrema s.l. com ácido salazínico 02
Diversidade das espécies de Parmotrema s.l. com ácido salazínico 05
Ácido salazínico 06
Material and Methods 11
Results and discussion 13
Key to the species of Parmotrema s.l. with salazinic acid 14
Parmotrema acanthifolhim (Pers.) Spielmann & Marcelli 18
Parmotrema albinatum (Moon, Kurok. & Kashiw.) Blanco, Crespo,
Divakar, Elix & Lumbsch 19
Parmotrema austrocetratum Elix & Johnston 20
Parmotrema austromaculatum Spielmann & Marcelli 21
Parmotrema bifidum Spielmann & Marcelli 22
Parmotrema cetratum (Ach.) Hale 23
Parmotrema clavulifenim (Rãs.) Streimann 25
Parmotrema concors (Kremp.) Spielmann & Marcelli 26
Parmotrema concarrens Hale 27
Parmotrema coralliforme (Hale) Hale 28
Parmotrema cristifenim (Taylor) Hale 29
Parmotrema cristobaliae (Ferraro & Elix) Blanco, Crespo,
Divakar, Elix & Lumbsch 33
Parmotrema delicatalum (Vainio) Hale 34
Parmotrema elixii Spielmann & Marcelli 35
Parmotrema enteroxanthum Hale 36
Parmotrema embescens (Stirton) Krog & Swinscow 37
Parmotrema eurysacum (Hue)Hale 38
Parmotrema expansum Hale 40
Parmotrema flavescens (Kremp.) Hale 41
Parmotrema foliolosiim (C.W. Dodge) Spielmann & Marcelli 42
Parmotrema granulare (Asahina) Spielmann & Marcelli 43
Parmotrema herrei (Zahlbr.) Spielmann & Marcelli 44
Parmotrema lacteum Marcelli & Spielmann 46
Parmotrema latissimum (Fée) Hale 47
Parmotrema leucosemothetum (Hue) Hale 48
Parmotrema lichexanthonicum Eliasaro & Adler 50
Parmotrema livido-tesselatiim (Hue) Spielmann & Marcelli 51
Parmotrema magmim (Lynge) Spielmann & Marcelli 52
Parmotrema mantiqaeirense Hale 53
Parmotrema margaritatum (Hue) Hale 54
Parmotrema masonii Ferraro 55
Parmotrema maximum (Hue) Spielmann & Marcelli 56
Parmotrema mirandum (Hale) Hale 58
Parmotrema neotropicum Kurok 59
Parmotrema nudiim (Hue) Spielmann & Marcelli 60
Parmotrema nylanderi (Lynge) Hale 61
Parmotrema parahypotropum (W.L. Culb.)Hale 62
Parmotrema param ore li ense W.L. Culb. & CF. Culb 63
Parmotrema pectinatum Jungbluth& Marcelli 64
Parmotrema permaculatum (Hale) Kurok 64
Parmotrema petropoliense ( Zahlbr.) Spielmann & Marcelli 65
Parmotrema pontagrossense (Eliasaro & Adler) Blanco, Crespo,
Divakar, Elix & Lumbsch 66
Parmotrema pseudoreticulatum (Tavares) Hale 67
Parmotrema radiatum (Lynge) Spielmann & Marcelli 68
Parmotrema ramusculum (Hale) Hale 69
Parmotrema reitzii Hale 70
Parmotrema reparatum (Stirton) Blanco, Crespo, Divakar, Elix & Lumbsch 71
Parmotrema reterimulosum (Zahlbr.) Spielmann & Marcelli 72
Parmotrema reticulatum (Taylor) M. Choisy 73
Parmotrema mm inatum (Zahlbr.) Blanco, Crespo, Divakar, Elix & Lumbsch 75
Parmotrema riiptum (Lynge) Hale ex DePriest & B. Hale 77
Parmotrema sieberi (C.W. Dodge) Spielmann & Marcelli 78
Parmotrema spinibarbe (Kurok.) Hale ex DePriest & B. Hale 79
Parmotrema stuppeum (Taylor) Hale 81
Parmotrema subcaperatum (Kremp.) Hale 82
Parmotrema subisidiosum (Mílll. Arg.) Hale 84
Parmotrema subsumptum (Nyl.)Hale 85
Parmotrema subtinctorium (Zahlbr.) Hale 87
Parmotrema ultralucens (Krog) Hale 88
Parmotrema verrucisetosum Sipman 89
Parmotrema warmingii (Vainio) Spielmann & Marcelli 90
Nomina inquirenda 93
Excluded name 97
Final remarks 98
References 99
Lista de tabelas
Tabela 1 . Cronologia das espécies de liquens parmotremóides com salazínico 07
Tabela 2. Espécies de Parmotrema s.l. com ácido salazínico citadas para América do Sul,
excetuando espécies brasileiras propostas por Canêz (2005), Spielmann (2005) e
Benatti(2005) 10
Tabela 3. Main characters of Parmotrema subcaperatum and P. embescens 83
Lista de Figuras
Figure 1 -Holotype of Parmotrema acanthifolium (P) 111
Figure 2 -Part of the holotype of Parmotrema albinatum (TNS) 111
Figure 3 - Part of the holotype of Parmotrema albinatum (TNS), underside 112
Figure 4 - Holotype of Parmotrema austrocetratum (CHR) 1 12
Figure 5 - Holotype of Parmotrema austrocetratum (CHR), upper surface with schizidia 113
Figure 6 - Holotype of Parmotrema austromaculatum (SP) 113
Figure 7 -Holotype of Parmotrema bifidum (S) 114
Figure 8 - Holotype of Parmotrema bifidum, undersurface 1 14
Figure 9 - Holotype of Parmotrema bifidum, furcated cilia 1 14
Figure 10 - Holotype of Parmotrema bifidum (S), upper surface with strong effigurate
maculae 115
Figure 1 1 - Lectotype of Parmotrema cetratum (H-ACH) 115
Figure 12 - Part of the lectotype of Parmotrema clavuliferum (H) 116
Figure 13 -Lectotype of Parmotrema concors (M) 116
Figure 14 - Part o f the holotype of Parmotrema concurrens (US) 117
Figure 15 - Part of the holotype of Parmotrema coralliforme (US), showing arbuscular
structures 117
Figure 16 - Lectotype of Parmotrema cristiferum (FH) 118
Figure 17 -Holotype oíParmelia mesotropa í. sorediosa (G) 118
Figure 18 -Holotype of Parmotrema cristobaliae (CTES) 119
Figure 19 -Holotype of Parmotrema delicatulum (TUR-V) 119
Figure 20 - Part of the isotype of Parmotrema e lixii (CANB) 120
Figure 21 -Isotype of Parmotrema enteroxanthum (UPS) 120
Figure 22 - Lectotype of Parmotrema erubescens (BM) 121
Figure 23 - Holotype of Parmotrema eurysacum (P) 122
Figure 24 - Part of the holotype of Parmotrema expansum (US) 123
Figure 25 - Holotype of Parmotrema flavescens (M) 123
Figure 26 - Holotype of Parmotrema foliolosum (FH) 124
Figure 27 - Holotype of Parmotrema foliolosum, showing the undersurface 1 24
Figure 28 - Holotype of Parmotrema granularis (TNS) 125
Figure 29 - Holotype of Parmotrema herrei (FH) 126
Figure 30 - Holotype of Parmotrema lacteum (SP) 126
Figure 31 -Lectotype of Parmotrema latissimum (G) 127
Figure 32 - Holotype of Parmotrema leucosemothetum (P) 127
Figure 33 - Holotype of P leucosemothetum, upper surface with strong effigurate
maculae 128
Figure 34 - Holotype of Parmotrema leucosemothetum, under marginal zone 128
Figure 35 - Holotype of Parmotrema lichexanthonicum (UPCB) 129
Figure 36 - Lectotype of Parmotrema livido-tesselatum (P) 129
Figure 37 -Holotype of Parmotrema magnum (S) 130
Figure 38 - Isotype of Parmotrema magnum (LD), showing the rugose and lacerate
apothecia 130
Figure 39 -Holotype oíParmelia microdactyla (UPS) 131
Figure 40 - Holotype of Parmotrema microdactyla, showing the lobules 131
Figure 41 -Isotype of Parmotrema mantiqueirense (US) 132
Figure 42 - Part of the lectotype of Parmotrema margaritatum (P) 132
Figure 43 -Holotype of Parmotrema masonii (CTES) 133
Figure 44 - Part o f the holotype of Parmotrema maximum (P) 133
Figure 45 -Partof the holotype ofParmoírema mirandum (US) 134
Figure 46 -Holotype ofParmotrema neotropicum (US) 134
Figure 47 -Partof the holotype of Parmo trema niidum (P) 135
Figure 48 -Holotype of Parmo trema nylanderi (S) 135
Figure 49 -Holotype of Parmo trema parahypotropnm (TNS) 136
Figure 50 - Holotype ofParmotrema parahypotropum, showing the soralia 1 36
Figure 51 -Holotype of Parmotrema paramoreliense (DUKE) 137
Figure 52 - Holotype of Parmotrema paramoreliense, showing effigurate maculae and the
black marginal line 137
Figure 53 -Partof the holotype of Parmotrema pectinatum (SP) 138
Figure 54 -Holotype of Parmotrema permaculatum (US) 138
Figure 55 -Partof the holotype of Parmotrema petropoliense (W) 139
Figure 56 - Holotype ofParmotrema petropoliense, showing cilia and soralia 139
Figure 57 -Holotype of Parmotrema pontagr o ssense (UPCB) 140
Figure 58 - Syntype of Parmotrema pseudoreticulatum (F) 140
Figure 59 - Lectotype of Parmotrema radiatum (S) 141
Figure 60 - Holotype ofParmotrema ramusculum (BM) 141
Figure 61 -Holotype ofParmotrema reitzii (US) 142
Figure 62 - Holotype ofParmotrema reitzii, showing the undersurface 1 42
Figure 63 - Lectotype ofParmotrema reparatum (BM) 143
Figure 64 - Holotype of Parmotrema reterimiãosiim (WU) 143
Figure 65 - Duplicate of the lectotype ofParmotrema reticulatum (BM) 1 44
Figure 66 - Lectotype ofParmotrema ruminatum (W) 144
Figure 67 - Holotype ofParmotrema niptum (S) 145
Figure 68 - Part of the holotype of Parmotrema sieberi (FH) 145
Figure 69 - Holotype of Parmotrema spinibarbe (TNS) 146
Figure 70 - Holotype of Parmotrema spinibarbe, showing the cespitose cilia 1 46
Figure 7 1 - Isotype of Parmotrema spinibarbe (G), showing the botryose structure 1 47
Figure 72 - Holotype of Parmotrema stuppeum (FH) 147
Figure 73 - Holotype of Parmotrema subcaperatum (M) 148
Figure 74 - Lectotype ofParmotrema subisidiosum (G) 148
Figure 75 - Holotype ofParmotrema subsumptum (H-NYL) 149
Figure 76 - Holotype ofParmotrema subtinctorium (WU) 149
Figure 77 -Holotype ofParmotrema ultralucens (O) 150
Figure 78 -Holotype ofParmotrema verrucisetosum (B) 150
Figure 79 - Holotype of Parmotrema verrucisetosum, showing ciliate warts 1 50
Figure 80 -Lectotype ofParmotrema warmingii (M) 151
Figure 81 -Lectotype of Parmelia badia (P) 152
Figure 82 -Holotype ofP arme line lia afrocetrata (B) 152
RESUMO
As espécies do género Parmotrema s.l. com ácido salazínico são revisadas, com base
especialmente nos tipos das espécies aceitas e de seus sinónimos. São fornecidas chave e
descrições para 61 espécies, incluindo sinonímia, química, distribuição e discussão das
afinidades taxonômicas de cada táxon.
Foi utilizada metodologia clássica em Liquenologia, contando com uso de microscópio óptico e
estéreo scópico, bem como ferramentas químicas para identificação das substâncias de
importância taxonômica, tais como testes de coloração, lâmpada UV, cromatografia em camada
delgada (TLC) e, para várias espécies, cromatografia líquida de alta eficiência (CLAE).
Duas espécies novas são descritas: Parmotrema anstromaculatum e P. bifidiim. Uma nova
combinação dentro de Parmelinella foi proposta, P. afrocetrata (Elix, Fischer & Killmann)
Marcelli & Spielmann, e 14 novas combinações em Parmotrema: P. acanthifolium (Pers.)
Spielmann & Marcelli, P. concors (Kremp.) Spielmann & Marcelli, P. folio lo sum (C.W. Dodge)
Spielmann & Marcelli, P. granularis (Asahina) Spielmann & Marcelli, P. herrei (Zahlbr.)
Spielmann & Marcelli, P. livido-tesselatum (Hue) Spielmann & Marcelli, P. magnum (Lynge)
Spielmann & Marcelli, P. maximum (Hue) Spielmann & Marcelli, P. nudum (Hue) Spielmann &
Marcelli, P. petropoliense ( Zahlbr.) Spielmann & Marcelli, P. radiatum (Lynge) Spielmann &
Marcelli, P. reterimulosum (Zahlbr.) Spielmann & Marcelli, P. sieberi (C.W. Dodge) Spielmann
& Marcelli e P. warmingii (Vainio) Spielmann & Marcelli. Doze táxons permanecem sem status
definido e estão dentro dos Nomina inquirenda.
ABSTRACT
Species of P armo trema s.l. with salazinic acid are revised, based on the types of the accepted
species as well as his synonyms. A key and descriptions for the 61 species are given, and the
synonymy, chemistry, distribution and taxonomic affinities of each species are discussed.
Classical methods in Lichenology were employed, using stereomicroscope and optical
microscopy for the observations, and spot tests, TLC and HPLC for the chemical analysis.
Two new species are described: Parmotrema austromaculatum e P. bifidum. One new
combination in Parmelinella was made, P. afrocetrata (Elix, Fischer & Killmann) Marcelli &
Spielmann, and 14 new combinations in Parmotrema: P. acanthifolhim (Pers.) Spielmann &
Marcelli, P. concors (Kremp.) Spielmann & Marcelli, P . foliolosiim (C.W. Dodge) Spielmann &
Marcelli, P. granularis (Asahina) Spielmann & Marcelli, P. herrei (Zahlbr.) Spielmann &
Marcelli, P. livido-tesselatiim (Hue) Spielmann & Marcelli, P. magnum (Lynge) Spielmann &
Marcelli, P. maximum (Hue) Spielmann & Marcelli, P. nudum (Hue) Spielmann & Marcelli, P.
petropoliense ( Zahlbr.) Spielmann & Marcelli, P. radiatum (Lynge) Spielmann & Marcelli, P.
reterimiãosiim (Zahlbr.) Spielmann & Marcelli, P. sieberi (C.W. Dodge) Spielmann & Marcelli
and P. angustatum (Vainio) Spielmann & Marcelli. Twelve taxa remain without definite status
and are included in the Nomina inquirenda.
Introdução
O levantamento de Parmeliaceae na região central do Rio grande do Sul (Spielmann 2005)
demonstrou que há um grupo cosmopolita de fungos liquenizados com problemas taxonômicos
interessantes e complexos formado por espécies dos géneros Canomaculina, Parmotrema e
Rimelia com ácido salazínico na medula.
Entre o período da defesa do mestrado e início do projeto de doutorado (em agosto de 2005),
foi proposta a sinonimização dos géneros Canomaciãina e Rimelia sob Parmotrema (Blanco et
ai. 2005), que são designados atualmente como parmotremóides ou Parmotrema s.l.
Os problemas taxonômicos e nomenclaturais englobam os tipos de todos os nomes existentes,
incluindo seus sinónimos. Assim, este é basicamente um estudo de materiais- tipo, às vezes
apoiado em coletas recentes, embora sem ter o objetivo de ser um trabalho ílorístico.
Histórico de Parmotrema s.l. com ácido salazínico
Os nomes novos, nomes inválidos, novas combinações e proposições de sinónimos estão
ordenados cronologicamente na Tabela 1 .
No breve histórico que se segue, a ênfase é colocada principalmente naqueles autores que
propuseram nomes ou combinações novas, bem como descreveram novas espécies. Não são
incluídos os trabalhos de flora, listagens ou que comentam espécies publicadas validamente em
outro local.
Parmotrema Massalongo
A história dos liquens parmotremóides inicia com a proposição do género Parmotrema
(Massalongo 1860). Neste trabalho, Massalongo utilizou como caráter primordial para a
separação genérica a presença de apotécios perfurados, os quais, dentro do então género
Parmelia, eram conhecidos em somente uma espécie, Parmelia perforata (Wulfen) Ach.
Contudo, Parmotrema não foi bem aceito. De fato, no primeiro grande trabalho sobre
Parmeliaceae brasileiras, Vainio (1890) usou o nome Parmelia, sequer citando o género
Parmotrema. Apesar disso, uma das seções que Vainio propôs dentro de Parmelia,
Amphigymnia, correspondia muito bem ao conceito de Parmotrema de Massalongo, e essa
correspondência apareceu assim listada por Zahlbruckner (1907).
Posteriormente, Dodge (1959) q\q\ ou Amphigymnia à categoria de subgênero, e foi sob esta
denominação que Hale (1965) efetuou uma monografia mundial do grupo.
Nesse meio tempo, somente um liquenólogo francês Choisy (1952) havia utilizado o nome
Parmotrema, porém em um senso diferente do que estava sendo usado pela literatura.
Alguns anos mais tarde, Hale (1974) restabeleceu o género Parmotrema, combinando nele
todas as espécies que havia incluído em sua monografia, além das espécies pertencentes à
Parmelia seção Irregulares, de que não havia tratado. Alguns liquenólogos demoraram a aceitar
esta divisão (por exemplo, Krog & Swinscow 1981), apesar de posteriormente terem mudado de
ideia (Krog & Swinscow 1983, Swinscow & Krog 1988). Hoje em dia o nome Parmotrema é
amplamente utilizado, embora com uma circunscrição um tanto obscura. Cerca de 300 espécies
são conhecidas para o mundo (Nash & Elix 2002b), das quais 94 foram citadas para o Brasil
(Marcelli 2008).
Canomaculina Elix & Hale e Rimeliella Kurok.
Elix & Hale (1987) propuseram o género Canomaculina para um grupo de espécies (o grupo
Parmelia pilosa) que tinham em comum (1) cílios robustos, marcadamente pontiagudos e
geralmente furcados, (2) superfície superior com máculas efiguradas e (3) conídios filiformes
com 12-16 um de comprimento, recombinando nele três espécies: Canomaculina consors (Nyl.)
Elix & Hale, C. muelleri (Vainio) Elix & Hale e C. pilosa (Stizenb.) Elix & Hale. Estas espécies
haviam sido tratadas por Hale (1974, 1976) dentro do género Parmelina Hale, caracterizado por
apresentar espécies com lobos estreitos, adnatos, (1,0) 2,0-4,0 ( _ 5,0) mm de largura, e pela
ausência de ácido úsnico no córtex superior.
Poucos anos depois, em um estudo bastante acurado, Kurokawa (1991) propôs o género
Rimeliella para acomodar sete espécies com lobos largos (5,0-15,0 mm) que estavam dentro de
Parmotrema^ e que apresentavam como características principais (1) rizinas dimórficas, (2)
superfície inferior de marrom a marrom-pálido (3) superfície superior com máculas efiguradas e
(4) cílios frequentemente pouco ramificados.
Entretanto, Elix (1997) afirmou que Canomaculina e Rimeliella exibiam uma sobreposição de
caracteres, diferindo apenas na largura dos lobos, e propôs a sinonimização de Rimeliella sob
Canomaculina.
Atualmente são conhecidas cerca de 20 espécies de Canomaculina (Hale & DePriest 1999),
das quais 15 são citadas para o Brasil (Marcelli 2008).
Rimelia Hale & Fletcher
O género Rimelia foi proposto por Hale & Fletcher (1990), para acomodar um grupo de espécies
(seção Irregulares Vainio), que compartilhavam (1) um córtex superior com máculas reticuladas,
(2) rizinas relativamente longas (até 2 mm) normalmente atingindo a margem inferior dos lobos,
não ou pouco ramificadas até esquarrosas, (3) apotécios perfurados e (4) conídios de bacilares a
filiformes com 9-16 um de comprimento. Destes caracteres, o mais importante era sem dúvida a
presença de máculas reticulares, já que existem espécies de Parmotrema com uma ou outra
dessas características.
Aproximadamente 20 espécies de Rimelia são conhecidas (Spielmann & Marcelli 2006).
Histórico das Espécies de Parmotrema s.l. com ácido salazínico
Acharius, o "Pai da Liquenologia" (Hale 1983), foi o primeiro (Acharius 1803) a propor uma
espécie parmotremóide com ácido salazínico: Parmelia perlata (Huds.) Ach. var. olivaria Ach.
Esta espécie já foi considerada como válida (Hale & Fletcher 1990), embora atualmente exista
uma proposta de rejeitar este nome em favor de Parmotrema pseudoreticulatum (Tavares) Hale
(Hawksworth et ai. 2003). A outra espécie proposta por Acharius (1814) é Parmelia cetrata Ach.
[Parmotrema cetratum (Ach.) Hale], com distribuição mundial e amplamente citada na
literatura. Sua variação morfológica é considerada grande, e por este motivo detém uma extensa
lista de sinónimos (Hale & Fletcher 1990).
Nesse meio tempo Lamarck & De Candolle (1805) descreveram Lichen perlatus a ciliatus,
considerado atualmente um sinónimo de Parmotrema reticulatum (Taylor) M. Choisy.
Elias Fries também desenvolveu trabalhos com liquens. Em seu Systema Orbis Vegetabilis
(Fries 1825) descreveu Parmelia polita Fr., hoje considerada sinónimo de Parmotrema cetratum.
Um ano depois, Persoon (in Gaudichaud-Beaupré 1826) descreveu três espécies, Parmelia
acanthifolia Pers., P. badia Pers. e P. maura Pers., as quais são consideradas sinónimos de
Parmotrema cetratum (Hale & Fletcher 1990).
Duby (1830) e Wallroth (1831) descreveram, respectivamente, Parmelia perlata g ciliata
(Lam. & DC.) Duby e Parmelia perlata var. tentacidata Wallr. Atualmente, ambas são
sinónimos deRimelia reticidata (Hale & Fletcher 1990).
Parmelia reticulata foi proposta por Taylor (in Mackay 1836). O caráter "reticulado" foi
adequadamente atribuído às máculas, e não às rachaduras do líquen ("the surface marked with
minute, whitish, elevated, reticulated Unes"). Este aspecto, contudo, viria causar muita confusão
no futuro, pois as máculas são geralmente difíceis de serem interpretadas (Brodo 1965). Em
outra publicação, Taylor (1847) descreveu P. cristifera eP. stuppea, ambas consideradas válidas
atualmente.
Aparentemente a primeira ilustração de um líquen parmotremóide com salazínico foi
publicada por Fée (1837), ao descrever Parmelia latíssima, espécie reconhecida até nossos dias.
Em seguida, Meyen & Flotow (1843) publicaram uma bela ilustração para Parmelia perforata
var. replicata (hoje sinónimo de Parmotrema cetratum), baseada em material proveniente do Rio
de Janeiro. Outra variedade proposta neste artigo, Parmelia perforata var. ulophylla, é hoje
sinónimo de Parmotrema cristiferum.
Em 1866, Lindsay considerou Parmelia reticulata de Taylor como uma variedade de
Parmelia laevigata, contudo esta proposição não teve aceitação (Hale & Fletcher 1990).
Krempelhuber (1869, 1873, 1876a, 1876b, 1878, 1881) publicou uma série de espécies e
variedades. De suas proposições, Parmelia glaberrima var. flavescens, Parmelia subcaperata e
Parmelia uruguensis são aceitas atualmente. Parmelia angustata, P. perforata var. corniculata e
P. perforata f integra são sinónimos de Parmotrema cetratum. Parmelia hildebrandtii é
sinónimo de Parmotrema cristiferum, e Parmelia concors é sinónimo de Parmotrema
reticulatum.
Somente uma das proposições novas de Nylander (1869, 1878, 1885, 1886, 1900), Parmelia
subsumpta, é aceita atualmente. Sua combinação de Parmelia flavescens passou para
Parmotrema. Parmelia praeperlata é sinónimo de Parmotrema reticulatum, e Parmelia
imperforata é sinónimo de Parmotrema subcaperatum.
Utilizando seu conhecimento dos conídios, Stirton (1877, 1877-1878), após discutir as
diferenças entre liquens de vários locais do mundo, propôs três espécies, duas das quais
(Parmelia erubescens e P. reparatá) são aceitas atualmente. Parmelia owaniana é considerada
sinónimo de Parmotrema cetratum.
Míiller Argoviensis (1880, 1883, 1886, 1891 a/b, 1892, 1894) também foi prolífico na
proposição de novos nomes ou novas combinações. Contudo, somente Parmelia cetrata var.
subisidiosa (Parmotrema subisidiosum) sobreviveu até os dias de hoje.
Surgiu então E. Vainio, o "Pai da Liquenologia Brasileira" (Marcelli & Ahti 1998). Suas
descrições são claras e detalhadas e seu esquema de classificação infragenérico teve um grande
impacto, formando a base do sistema atual (Feuerer 1998). De suas espécies e variedades novas
(Vainio 1890, 1909a/b, 1918), Parmelia delicatula resistiu ao tempo.
Em 1896 Harmand descreveu Parmelia perforata var. claudelii e Vainio (1909a) recombinou-
a como P. claudelii. Atualmente, é considerada sinónimo de Parmotrema stuppeum.
Descrições e comentários detalhados foram publicados por Hue (1899), sendo Parmelia
eurysaca, P. leucosemotheta eP. margaritata espécies válidas até hoje.
Em suas contribuições, Zahlbruckner (1902, 1904, 1908, 1909, 1926, 1928, 1930b) continuou
a tradição iniciada por Vainio de fornecer descrições completas e acuradas sobre os liquens, além
de uma listagem completa das espécies conhecidas (Zahlbruckner 1930a, 1934). Porém, somente
Parmelia subtinctoria, P. ramescens e P. ruminata continuam sendo usadas (estas últimas
espécies publicadas em Magnusson & Zahlbruckner 1944).
Jatta (1909) publicou Parmelia urceolata f ciliata, atualmente sinónimo de Parmotrema
reticulatum. Deve-se notar, porém, que o nome que aparece na lista de sinónimos de Hale &
Fletcher (1990) para Rimelia reticulata, é Parmelia perlata var. ciliata.
Continuando os bons modelos de descrição apresentados por Vainio, Hue e Zahlbruckner,
Lynge (1914, 1924), ao estudar as Parmelia coletadas por Malme na América do Sul, descreveu
4
Parmelia nylanderi e P. rupta, válidas até hoje. Parmelia magna e P. radians são consideradas
sinónimos de Parmotrema delicatulum (Hale 1965). Parmelia cetrata subsp. radiata Lynge está
na sinonímia de Parmotrema cetratum e Parmelia hieronymi é sinónimo de Parmotrema
uruguense.
Ao descrever os liquens do México coletados pelo Irmão Arséne, Bouly de Lesdain (1914)
propôs Parmelia moreliensis, espécie aceita até hoje.
Fink (in Fink & Fuson 1919) propôs Parmelia ciliata, uma combinação supérflua, que já
havia sido publicada por Nylander (1878), e que hoje está na sinonímia de Parmotrema
reticulatum.
Por essa época, a aplicação de nomes iguais a espécies certamente diferentes era algo comum
e confuso na literatura. Du Rietz (1924a) contribuiu bastante ao esclarecer a confusão que existia
em torno de Parmelia perforata. Sua combinação Parmelia trichothera var. claudelii, contudo
(Du Rietz 1924b), não é reconhecida atualmente (sinónimo de Parmotrema stuppeum).
Gyelnik (1931, 1934) contribuiu para o estudo de Parmotrema s.L Das suas quatro
proposições, nenhuma permanece atualmente, porém seu entendimento dos liquens e dos
procedimentos nomenclaturais era bastante avançado para o seu tempo (Hale 1990).
Cengia Sambo (1930, 1939a/b/c) trabalhou com material proveniente do Brasil {Parmelia
lencoxantha f. firma), Argentina e Etiópia, para o qual propôs três táxons intra-específicos,
nenhum dos quais aceitos hoje em dia.
Em seus estudos sobre liquens de Taiwan (antiga Formosa), Asahina (1940)descreveu
Parmelia cetrata f. granularis. Posteriormente (Asahina 1952), sinonimizou este táxon com
Parmelia cetrata f. subisidiosa. Hoje é considerado sinónimo de Parmotrema reticulatum.
Parmelia clavulifera, proposta por Rásánen (1944) já foi considerada uma boa espécie
(Kurokawa 1991, Kurokawa & Lai 2001, Moon et ai. 2001) ou sinónimo de Parmotrema
reticulatum (Hale & Fletcher 1990, Divakar et ai. 2005). As outras duas proposições de Rásánen
(19 '47), Parmelia perforata var. microspora eP. cinerascens var. saxicola, são hoje sinónimos
de Parmotrema cetratum (Hale & Fletcher 1990).
Magnusson (in Magnusson & Zahlbruckner 1944) publicou somente um táxon no grupo,
Parmelia hawaiiense, que é aceito atualmente.
Outro táxon discutido, recentemente estudado sob o prisma da biologia molecular (Divakar et
ai. 2005) é Parmelia pseudoreticulata, descrita por Tavares (1945).
Dodge (1959) publicou várias espécies novas ou elevou para o nível de espécie variedades ou
formas propostas por Hue e por Muller Argoviensis. Destas suas combinações, somente
Parmelia subisidiosa (hoje Parmotrema) é aceita.
Hale (1960, 1965, 1971, 1977, 1990) inicia, então, uma série de estudos dos liquens
parmotremóides, especialmente das espécies tropicais, com uma extensa revisão de nomes
antigos e proposição de várias espécies novas, todas aceitas atualmente à exceção de Parmelia
microdactyla. O trabalho de Hale é de fundamental importância para qualquer estudo neste
grupo, visto que ele teve acesso aos tipos e a uma grande quantidade de material adicional,
proveniente do mundo inteiro.
Já por estes anos o estudo químico dos liquens contava com uma nova ferramenta: a
cromatografia em camada delgada. Fazendo uso desta técnica, e com propostas sobre especiação
em liquens, Culberson (1973) e Culberson & Culberson (1981) propuseram Parmelia
parahypotropa e Parmotrema paramoreliense, ambas boas espécies.
Outro importante estudioso do grupo foi Kurokawa, que fornece dados acurados sobre os
liquens que descreveu (Kurokawa 1974, 2001; Moon et ai. 2001), o que se reílete na aceitação
de todas suas espécies.
A combinação de características morfológicas e químicas sempre foi decisiva na aceitação de
espécies de Parmotrema. Quando a substância liquexantona foi detectada pela primeira vez em
liquens parmotremóides, ela justificou a descrição de Parmelia ultralucens (Krog 1974), uma
espécie com isídios. Posteriormente, esta substância foi descoberta em uma espécie sem isídios,
Parmotrema lichexanthonicum (Eliasaro & Adler 1997), e finalmente em uma espécie sorediada,
Parmotrema pontagrossensis (Eliasaro & Adler 1998).
Outras vezes, um caráter morfológico aberrante também suscita a suspeita de um novo táxon,
como ocorreu com Parmotrema arteagum, que Egan (1982) descreveu para o México. Esta
espécie tem conídios muito mais longos (16-23 um) que o usual em Parmotrema.
Até o final dos anos 1970, nenhuma espécie havia sido descrita por um liquenólogo latino-
americano. Iniciaram então os trabalhos de liquenólogo s argentinos: Ferraro (1979) propôs
Parmotrema masonii, Ferraro & Elix (1993, 2000) adicionaram duas novas espécies a esse país:
Rimelia cristobaliae e Canomacalina laciniella.
Adler (1988) realizou a primeira tese de doutorado com Parmeliaceae na América do Sul,
publicando Canomaculina tandilense e C. ventanica dentro do grupo (Adler & Elix 1987).
Elix publicou diversos estudos de espécies geralmente australianas ou sul-americanas (Adler
& Elix 1987; Elix & Johnston 1988; Ferraro & Elix 1993, 2000; Elix et ai 2002, 2005; Benatti et
ai. 2008), nos quais um dos pontos fortes é a informação química dada a cada espécie.
Estudando liquens das Guianas e arredores, Sipman & van Aubel (1992) descreveram
Parmotrema verrucisetosum, uma espécie única por apresentar verrugas ciliadas na superfície do
talo. Sipman (2005) teve também um papel importante ao disponibilizar publicamente uma nova
versão de uma chave para identificação que Hale havia feito para o neotrópico antes de seu
falecimento.
Na década de 1990, os brasileiros começaram a desenvolver estudos aprofundados em liquens
parmotremóides. Fleig (1997) efetuou uma revisão de Parmotrema, Rimelia e Rimeliella do Rio
Grande do Sul, tendo analisado uma grande quantidade de material e visto praticamente todos os
tipos disponíveis.
Ribeiro (1998) defendeu dissertação de mestrado sobre Parmeliaceae de regiões montanhosas
de São Paulo, Rio de Janeiro e Minas Gerais. Dentro do grupo em estudo, foram descritas
Parmotrema lobuliferum eP. neosiibcrinitam (Marcelli & Ribeiro 2002).
Como resultado de sua tese de doutorado, Eliasaro (2001) publicou duas espécies novas
contendo liquexantona (ver acima).
Nos anos recentes, quatro dissertações de mestrado (Canêz 2005, Spielmann 2005, Benatti
2005 e Jungbluth 2006) sobre Parmeliaceae, apresentaram 17 novas espécies de liquens
parmotremóides com salazínico, das quais Parmotrema anchietanum Marcelli, Benatti & Elix,P.
asperum Benatti, Marcelli & Elix, P. fleigiae Canêz & Marcelli, P. hypermaculatum Marcelli &
Benatti, P. lacteum Marcelli & Spielmann e P. pectinatum Jungbluth & Marcelli já foram
publicadas (Marcelli et ai. 2007, Benatii et ai. 2008). As 1 1 espécies restantes estão em fase de
publicação.
Assim, atualmente são conhecidas, dentro dos liquens parmotremóides com salazínico:
63 espécies aceitas pela literatura corrente e validamente publicadas;
1 1 espécies em fase de publicação;
120 nomes, representados pelos tipos das espécies aceitas e seus sinónimos;
39 espécies citadas para a América do Sul;
3 1 citadas para o Brasil;
Diversidade das espécies de Parmotrema s.l. com ácido salazínico
Excetuando as espécies recentemente descobertas por Canêz (2005), Spielmann (2005) e Benatti
(2005), que estão em fase de publicação, e comparando a distribuição das demais espécies,
percebe- se que a América do Sul é provavelmente o centro de diversidade deste grupo, com 39
espécies registradas (Tabela 2). Para o Brasil há 3 1 registros, mais do que o dobro em relação ao
segundo país, Venezuela, com 16 espécies citadas.
Dentro do Brasil, 13 espécies pertencentes a este grupo são citadas para o Paraná (Eliasaro
2006) 15 para São Paulo (Spielmann, dados não publicados) e 17 para o Rio Grande do Sul
(Spielmann 2006).
Fora da América do Sul ocorrem outras 16 espécies de Parmotrema, assim distribuídas:
América do Norte: P. arteagum, P. coralliforme, P. moreliense e P. permaculatum (Egan
1982, Hale 1977, Culberson & Culberson 1981, Kurokawa 2001, respectivamente).
Ásia, América do Norte, América Central, Europa e África: P. stuppeum (Hale 1965).
Ásia: P . parahypotropnm eP. ramusculum (Culberson 1973, Hale 1965).
Ásia e Oceania: P. pustulatum (Elix et ai 2002), veja porém P. elixii, adiante neste trabalho.
Ásia, Austrália e Havaí: P. hawaiiense (Hale & Fletcher 1990, Moon et ai. 2001).
Havaí: P. albinatum,P. ramescens eP. ruminatum (Moon et aí. 2001).
Oceania: P. ans troce tratum eP. embescens (Elix 1997, Hale & Fletcher 1990).
Europa: P. psendoreticulatum (Divakar et ai. 2005, Hale & Fletcher 1990).
Ácido salazínico
O primeiro registro deste ácido foi feito por Zopf em 1897, ao nomear o ácido que encontrou em
Stereocaalon salazinum Bory. Contudo, a substância que Zopf encontrou nesta espécie na
verdade tratava-se de ácido norstíctico (Asahina & Shibata 1954).
Asahina, em 1912, conseguiu isolar uma grande quantidade de ácido salazínico em Parmelia
cetrata Ach. (Asahina & Shibata 1954). O reconhecimento desta espécie como portadora de
ácido salazínico foi muito importante, já que ela teria uma ampla distribuição mundial (Hale &
Fletcher 1990) e é, juntamente com Parmotrema reticulatum, frequentemente citada como fonte
de ácido salazínico para substância padrão na comparação cromato gráfica em vários manuais
sobre liquens (p. ex. White & James 1985, Orange et ai. 2001).
O ácido salazínico pertence ao grupo das depsidonas e pode ser reconhecido nos testes de
coloração ("spot tests"), pela reação K+ amarelo -^ vermelho- sangue, C- e P+ amarelo forte
(Hale 1979, Orange et ai. 2001). Já na Cromatografia em Camada Delgada (CCD), o ácido
salazínico exibe um Rf relativo de 4 no solvente C e 10 no solvente A, formando pontos
alaranjados na placa (Orange et ai. 2001). Finalmente, na micro cristalização, usando o reagente
G.A.o-T., o ácido salazínico se micro cristaliza na forma de pequenos "barcos" amarelados
(Taylor 1967, Hale 1979).
Tabela 1. Cronologia das espécies de liquens parmotremóides com salazínico.
Ano
Nome original
Nome atualmente aceito
(P. = Parmotrema)
1803
P armelia perlata var. olivaria Ach.
P. pseudoreticulatum
1805
Lichen perlatus a ciliatiis Lam. & DC
P. reticulatum
1814
P armelia cetrata Ach.
P. cetratum
1825
P armelia polita Fr.
P. cetratum
1826
P armelia acanthifolia Pers.
P. cetratum
P armelia badia Pers.
P armelia maura Pers.
1830
P armelia perlata g ciliata (Lam. & DC) Duby
P. reticulatum
1831
P armelia perlata var. tentaculata Wallr.
P. reticulatum
1836
P armelia reticulata Taylor
P. reticulatum
1837
P armelia latíssima Fée
P. latissimum
1843
P armelia perforata var. ulophylla Mey. & Flot.
P. cristiferum
P armelia perforata var. replicata Meyen & Flotow
P. cetratum
1847
P armelia cristifera Taylor
P. cristiferum
P armelia stuppea Taylor
P. stuppeum
1866
P armelia laevigata var. reticulata (Taylor) Lindsay
P. reticulatum
1869
P armelia glaberrima var. flavescens Kremp.
P. flavescens
P armelia subsumpta Nyl.
P. subsumptum
1873
P armelia angustata Kremp.
P. cetratum
P armelia perforata var. corniculata Kremp.
1874
P armelia subcaperata Kremp.
P. subcaperatum
1876
P armelia glaberrima f. flavescens (Kremp.) Kremp.
P. flavescens
P armelia perforata f. integra Kremp.
P. cetratum
1877
P armelia hildebrandtii Kremp.
P. cristiferum
P armelia owaniana Stirton
P. cetratum
1877
1878
P armelia erubescens Stirton
P. erubescens
P armelia reparata Stirton
P. reparatum
1878
P armelia uruguensis Kremp.
P. uruguense
P armelia ciliata (Lam. & DC.) Nyl.
P. reticulatum
1880
P armelia latíssima f. flavescens (Kremp.) Miill. Arg.
P. flavescens
P armelia urceolata Eschw. var. nuda Miill. Arg.
P. subsumptum
1881
P armelia concors Kremp.
P. reticulatum
1883
P armelia urceolata var. subcetrata Miill. Arg.
P. reticulatum
1885
P armelia flavescens (Kremp.) Nyl.
P. flavescens
1886
P armelia praeperlata Nyl.
P. reticulatum
P armelia virens Miill. Arg.
P. reparatum
1890
P armelia delicatula Vainio
P. delicatulum
P armelia cetrata f. sorediifera Vainio
P. reticulatum
P armelia warmingii Vainio
P. cetratum
1891
P armelia hildebrandtii Kremp. f. sorediosa Miill. Arg.
P. cristiferum
P armelia mesotropa Miill. Arg. f. sorediosa Miill. Arg.
P armelia cetrata var. corniculata (Kremp.) Miill. Arg.
P. cetratum
1892
P armelia virens Miill. Arg. f. isidiosa Miill. Arg.
P. subtinctorium
1894
P armelia cetrata var. subisidiosa Miill. Arg.
P. subisidiosum
1896
P armelia perforata var. claudelii Harm.
P. stuppeum
1899
P armelia acanthifolia f. ornata Hue
P. reticulatum
P armelia eurysaca Hue
P. eurysacum
P armelia ghattensis Hue
P. reticulatum
P armelia hypotropa var. imperialis Hue
P. subsumptum
Parmelia imperforata Nyl.
P. subcaperatum
Parmelia latíssima Fée f. cristifera (Taylor) Hue
P. cristiferum
Parmelia leiícosemotheta Hue
P. leucosemothetum
Parmelia leiícosemotheta Hue f. isidiata Hue
P. subtinctorium
Parmelia livido-tessellata Hue
P. reticulatum
Parmelia margaritata Hue
P. margaritatum
Parmelia mauriensis Hue
P. flavescens
Parmelia máxima Hue
P. stuppeum
Parmelia mutata f. decorata Hue
P. reticulatum
Parmelia olivaria (Ach.) Hue
P. pseudoreticulatum
Parmelia reticulata f. nuda Hue
P. reticulatum
1902
Parmelia petropoliensis Zahlbr.
P. subsumptum
1904
Parmelia subcaperata f. ciliata Zahlbr.
P. delicatulum
1906
Parmelia herrei Zahlbr.
P. cetratum
1908
Parmelia dusenii Zahlbr.
P. uruguense
1909
Parmelia claudelii (Harm.) Vainio
P. stuppeum
Parmelia claudelii (Harm.) Vainio var. clemensiae Vainio
P. cristiferum
Parmelia urceolata f. ciliata (DC) Jatta
P. reticulatum
Parmelia protoflavescens Zahlbr.
P. flavescens
Parmelia pseudoflavescens Zahlbr.
P. flavescens
1914
Parmelia cetrata subsp. radiata Lynge
P. cetratum
Parmelia magna Lynge
P. delicatulum
Parmelia moreliensis B. de Lesd.
P. moreliense
Parmelia nylanderi Lynge
P. nylanderi
Parmelia radians Lynge
P. delicatulum
Parmelia rupta Lynge
P. ruptum
1918
Parmelia cetrata f. platyloba Vainio
P. reticulatum
1918
Parmelia ciliata (Lam. & DC) Fink
P. reticulatum
1924
Parmelia trichotera Hue var. claudelii (Harm.) Du Rietz
P. stuppeum
1924
Parmelia hieronymi Lynge
P. uruguense
1926
Parmelia reterimulosa J. Steiner & Zahlbr.
P. reticulatum
1928
Parmelia velutina Zahlbr.
P. subtinctorium
1930
Parmelia corrugis (Fr.) Miill. Arg. var. imperialis (Hue) Zahlbr.
P. subsumptum
Parmelia perforata var. ciliata Sambo
P. uruguense
Parmelia subtinctoria Zahlbr.
P. subtinctorium
1931
Parmelia protovirens Gyelnik
P. subtinctorium
Parmelia submesotropa Gyelnik
P. cristiferum
1934
Parmelia amphygymnioides Gyelnik
P. reticulatum
Parmelia cristifera Taylor f. cinerata Zahlbr.
P. cristiferum
Parmelia diffusoides Gyelnik
P. reticulatum
1939
Parmelia leucoxantha Miill. Arg. f. firma Sambo
P. subsumptum
Parmelia odontata var. rubiginosa Sambo
P. reticulatum
1940
Parmelia cetrata f. granularis Asahina
P. reticulatum
1944
Parmelia cetrata var. ulcerata Zahlbr.
P. reticulatum
Parmelia clavulifera Rásánen
P. clavuliferum
Parmelia hawaiiensis H. Magn.
P. hawaiiense
Parmelia ramescens Zahlbr.
P. ramescens
Parmelia ruminata Zahlbr.
P. ruminatum
1945
Parmelia pseudoreticulata Tavares
P. pseudoreticulatum
1947
Parmelia cinerascens var. saxicola Rásánen
P. cetratum
Parmelia perforata var. microspora Rásánen
1959
Parmelia decorata (Hue) C.W. Dodge
P. reticulatum
Parmelia foliolosa C.W. Dodge
P. reticulatum
Parmelia gossweileri C.W. Dodge
P. cristiferum
Parmelia imerinensis C.W. Dodge
P. cristiferum
Parmelia menziesii C.W. Dodge
P. cetratum
Parmelia ornata (Hue) C.W. Dodge
P. reticulatum
Parmelia sanctae-helenae C.W. Dodge
P. reticulatum
Parmelia sieberi C.W. Dodge
P. cristiferum
Parmelia subisidiosa (Miill. Arg.) C.W. Dodge
P. subisidiosum
1960
Parmelia microdactyla Hale
P. delicatulum
1965
Parmelia coralliformis Hale
P. coralliforme
Parmelia miranda Hale
P. mirandum
Parmelia ramuscula Hale
P. ramusculum
1971
Parmelia permacul ata Hale
P. permaculatum
1973
P arme lia parahypotropa W.L. Culb.
P. parahypotropum
1974
Parmelia spinibarbis Kurok.
P. spinibarbe
Parmelia ultralucens Krog
P. ultralucens
1977
Parmotrema concurrens Hale
Parmotrema conferendum Hale
Parmotrema enteroxanthum Hale
Parmotrema expansum Hale
Parmotrema neotropicum Kurok.
Parmotrema reitzii Hale
1979
Parmotrema masonii Ferraro
1981
Parmotrema paramoreliense W.L. Culb. & CF. Culb.
1982
Parmotrema arteagum Egan
1987
Canomaculina tandilensis Adler & Elix
P. tandilense
Canomaculina ventanica Adler & Elix
P. ventanicum
1988
Parmotrema austrocetratum Elix & Johnston
1990
Parmotrema mantiqueirense Hale
1992
Parmotrema verrucisetosum Sipman
1993
Rimeliella cristobaliae Ferraro & Elix
P. cristobaliae
1997
Parmotrema lichexanthonicum Eliasaro & Adler
1998
Rimelia pontagrossensis Eliasaro & Adler
P. pontagrossense
2000
Canomaculina laciniella Ferraro & Elix
P. laciniellum
2001
Parmotrema acutatum Kurok.
Parmotrema despectum Kurok.
Rimelia albinata K.H. Moon, Kurok. & Kashiw.
P. albinatum
2002
Canomaculina lobulifera Marcelli & Ribeiro
P. lobuliferum
Parmotrema neosubcrinitum Ribeiro & Marcelli
Rimelia pustul ata Elix & Bawingan
P. pustul atum
2005
Parmotrema afrocetratum Elix, Eb. Fischer & Killmann
2007
Parmotrema fleigiae Canêz & Marcelli
Parmotrema lacteum Marcelli & Spielmann
Parmotrema pectinatum Jungbluth & Marcelli
2008
Parmotrema anchietanum Benatti, Marcelli & Elix
Parmotrema asperum Benatti, Marcelli & Elix
Parmotrema hypermaculatum Benatti, Marcelli & Elix
10
Tabela 2. Espécies de Parmotrema s.l. com ácido salazínico citadas para América do Sul,
excetuando espécies brasileiras propostas por Canêz (2005), Spielmann (2005) e Benatti (2005).
Espécies
AR
BO
BR
CH
CO
EQ
GU
GF
PA
PE
SU
UR
VE
P. cetratum
+
+
+
+
+
+
+
+
P. clavuliferum
+
P. concurrens
+
P. conferenciam
+
+
+
+
P. cristiferum
+
+
+
+
+
+
P. cristobaliae
+
+
+
P. delicatulum
+
+
+
+
P. enteroxantham
+
P. eurysacum
+
+
+
P. expansum
+
P.fíavescens
+
+
+
P. laciniellum
+
P. latissimum
+
+
+
P. lenços emothetum
+
+
P. lichexanthonicnm
+
P. lobulifenim
+
P. mantiqiieirense
+
P. margaritatum
+
+
P. masonii
+
+
+
P. mirandum
+
+
P. neosiibcrinitum
+
P. neotropicum
+
+
P. nylanderi
+
P . paramoreliense
+
P. pontagrossense
+
P. reitzii
+
+
P. repara tum
+
P. reticulatum
+
+
+
+
+
P. raptam
+
+
+
P. spinibarbe
+
P. subcaperatum
+
+
+
+
+
P. subisidiosum
+
+
P. sabsamptam
+
+
+
P. subtinctorium
+
P. tandilense
+
+
P. ultralucens
+
+
+
+
P. uruguense
+
+
+
P. ventanicam
+
P. verracise tosam
+
+
TOTAL
14
02
31
02
04
02
02
06
03
01
10
16
Abreviações - AR: Argentina, BO: Bolívia, BR: Brasil, CH: Chile, CO: Colômbia, EQ: Equador, GU:
Guiana, GF: Guiana Francesa, PA: Paraguai, PE Peru, SU: Suriname, VE: Venezuela, UR: Uruguai.
11
Material and Methods
Specimens were described using standard stereoscopic and light microscopic examination.
Anatomical sections of apothecia and pycnidia were made with razor blades from hand.
Measurements were recorded from water mounts. Permanent slides were made for ali types
examined, using resin PVLG (8,33 g of polyvinyl alcohol, 50 ml of lactic acid, 50 ml of distilled
water and 5 ml of glycerin), following Morton et ai. (1993), adding cotton blue instead of
Melzer's reagent, and adaptation introduced by Dr. Rosely A.P. Grandi (pers. comm.) to study
anamorphic fungi in the Instituto de Botânica, SP, Brazil.
Spot tests were performed with potassium hydroxide (K), sodium hypochlorite (C), and para-
phenylenediamine (P) for ali specimens, except the oligotypes or when chemical examination
was not allowed by the herbarium curator. Ali taxa were also examined under UV light. Lichen
substances were mainly identified by J.A. Elix (Canberra, Austrália), from fragments of the types
sent to him, through TLC in solvent C and HPLC, following standard methods (Huneck &
Yoshimura 1996, Bungartz 2001, Orange et ai. 2001). Otherwise some published data based on
the types studied were also included, if considered reliable.
Ali specimens studied were also scanned at high resolution (1200 dpi), the resulting images
being part of a data bank. Minute important features were also photographed coupling a digital
camera to a stereomicroscope using a tripod.
The protocol used to describe the species (see below) was an adaptation of that used by the
GEL (Grupo de Estudos Liquenológicos) of Instituto de Botânica (see Canêz & Marcelli 2006,
Marcelli et ai. 2007). Two terms used to describe the maculation of the upper surface are here
being employed for the first time:
Hypermaculate: when the whitish macular regions are very abundant and cover mo st of the
surface; the algae free parts are by far more extensive than the green ones, rendering the
thallus a milky-grey color.
Pseiido-reticalar maculae: this pattern is formed by the confluence of punctiform or
effigurate maculae, giving as result a "reticular pattern" in the upper surface. However,
reticular maculae usually give rise to reticular cracks, while in pseudo -reticulate species
the cracks have usually an independent origin from the maculae.
A third term, arbusculum (pi. arbuscula), was used to the structures named as "coralloid
isidia" by Hale (1965), since they are different from anything that could be called "isidia" in
another species.
The curators of the following herbaria kindly loaned specimens (acronyms follow Index
Herbariorum): ASU, B, BAFC, BM, C, CANB, CHR, COLO, CTES, DUKE, F, FI, G, H, K, L,
LD, M, MIN, MSC, O, P, RO, S, SP, TNS, TUR, UPS, UPCB, US, W, and WU. The herbarium
FH do not sent specimens, but made available an opportunity to stay two weeks studying
specimens at Farlow. A two-week stay was performed in F, and four days in ICN.
Protocol to description of Parmotrema s.l. with salazinic acid
Species Author
Publication not abbreviated, vol. pp. year.
Basionym Author, Publication.
Type: Localization more complete as possible, leg. Xxxx (type of type: herbarium!; type of type:
herbarium!).
! = seen (one sign for each species)
= nomenclatural (homotypic) synonym
= taxonomic (heterotypic) synonym
12
Type description
Thallus color, laciniate/ lobate, adnate/ loosely adnate, substrate, xx cm broad. Laciniael Lobes
dichotomous/ irregularly branched, contiguous/ laterally overlapped/ crowded, xxmm wide,
surface continuous/ reticulate/ cracked, smooth/ scrobiculate/ rugose, lustrous/ sublustrous/
opaque, becoming rugose and cracked in/towards the center; apical zone truncate/ subtruncate/
rounded; margin smooth/ crenate/ crenulated/ incised/ irregular/ erose/ sublacinulate, plane/
undulated. Maculae absent or weak/ distinct/ strong/ extensive (hypermaculate), punctiform/
effigurate/ reticular/ pseudo-reticular/ irregular, laminai, marginal, just in the amphithecium,
originating cracks/ pseudo cyphellae/ soredia/ pustules. Cilia black/ absent/ color, simple/
squarrose/ furcated/ irregularly branched /dichotomous /cespitose, xx-xxmm, contiguous/
abundant/ frequent/ few/ rare, present only in the axils. Lobiãesl Phyllidial Dactyls. Lacinules
absent or simple/ branched, present in the center/ in the whole thallus, plane/ canaliculated, apex
rounded/ truncate/ acute, xx-xx x xx-xx mm. Pustules absent or rugose/ capitate, distribution,
originating soredia/ remaining entire. Soralia absent or color, type, linear continuous/ linear
interrupted/ capitate/ extensive/ labriform/ orbicular, laminai/ marginal/ submarginal/ on the
lacinules ápices; soredia farino se/ granular/ isidioid. Isidia absent or base color, apex color,
simple/ branched/ coralloid, xx-xx x xx-xx mm, erect/ straight/ procumbent, firm/ friable, apex
ciliate/ sorediate, laminai/ marginal/ on the lacinules ápices, originating... . Medulla white/
yellow/ ochre/ brown, color proportion. Undersurface color, lustrous/ opaque/ shiny, smooth/
rugose/ papillate/ veined, with/ without cracks; marginal zone absent or color, opaque/ lustrous/
shiny, xx mm wide, naked/ rhizinate, with sharp/ attenuated limit, smooth/ rugose/ papillate/
veined; rhizines black/ color/ concolor to the undersurface, simple/ furcated/ irregularly
branched/ dichotomous/ squarrose, xx x xxmm, few/ frequent/ abundant/ tomentose, evenly
distributed/ irregularly distributed/ distributed in groups.
Apothecia absent or plane/ concave/ cupuliform/ convex/ urceolate, xxmm in diameter,
sessile/ stipitate, laminai/ submarginal/ on the laciniae ápices, margin smooth/ crenate/ dentate/
coronate/ ciliate/ pustulate/ isidiate/ lobulate/ sorediate, amphithecium maculate/ rugose/ isidiate/
sorediate/ smooth/ ciliate, disc color, epruinose/ pruinose, perforate/ imperforate; ascospores
ellipsoid/ oval/ globo se/ oblong, xx-xx x xx-xx um, episporium xx um. Pycnidia absent or
laminai/ submarginal/ apical, conspicuous/ inconspicuous, with/ without prominent margin,
abundant/ frequent/ few/ rare, ostiole black/ brown; conidia filiform/ sublageniform/ bifusiform/
unciform/ bacilliform, xx-xx x ca. xx um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry:
Remarks:
About the species (main features, cio se taxa, etc.)
Taxonomic (history of the taxon and its taxonomic problems; opinions from several authors;
explanation of our decision; excluded synonyms).
The type (s) (full comments about the size, number of parts, preservation, not seed features,
annotations by others, etc.)
Distribution: verify the synonyms!
Additional specimens examined:
13
Results and discussion
In the following account, 61 species of Parmotr ema s.l. with salazinic acid are keyed out,
described, commented and illustrated. Due to time constrains, 12 species were not covered, but
they will be included in the text submitted to publication. They are: Parmotrema aciitatam
Kurok., P. anchietanum Marcelli, Benatti & Elix, P. asperum Benatti, Marcelli & Elix, P.
arteagum Egan, P. despectum Kurok., P. fleigiae Canêz & Marcelli, P. hypermacalatiim
Marcelli & Benatti, P. laciniellum (Ferraro & Elix) Blanco, Crespo, Divakar, Elix & Lumbsch,
P. neosubcrinitum Ribeiro & Marcelli, P. tandilense (Adler & Elix) Blanco, Crespo, Divakar,
Elix & Lumbsch, P. uruguense (Kremp.) Hale and P. ventanicum (Adler & Elix) Blanco,
Crespo, Divakar, Elix & Lumbsch.
Some species features, although already employed by some authors in a greater or lesser
extension, were fully considered here, as the conidial type and size, the development and
morphology of soredia and isidia, the type and intensity of the maculation, the cilia length, width
and ramification, the presence/absence of certain chemical substances, the underside surface and
the type if ramification of rhizines.
The application of these characters, fully described in the types studied, resulted in the
proposition of two new species, 14 new combinations in Parmotrema, and one new combination
inP arme Une lia. Due to several factors, 12 taxa remain in the Nomina inquirenda.
It was realized that the present species concept in the Parmotrema with salazinic acid is
unsatisfactory, since several well-defined species were "hidden" under long lists of synonyms,
usually accepted uncritically ali over the world. Even trying to use a narrow species concept,
some special problems remain unsolved, like the present concept of Parmotrema cetratum and P.
reticulatum. Since this study is primarily based on the types available, additional specimens can
help to understand better the specific limits in these species, providing new data relating to the
distributional patterns, together with chemical and morphological features.
14
Key to the species oiParmotrema s.l. with salazinic acid
1 Upper córtex yello w or y ellowish in herbarium; usnic acid evidently present 2
1 Upper córtex gray or brownish in herbarium; usnic acid, when present, revealed only
through chemical analysis 10
2 (1) Thallus with isidia, soredia or pustules 3
2 Thallus without isidia, soredia or pustules 7
3 (2) Thallus with isidia 4
3 Thallus with soredia or pustules 5
4 (3) Maculae inconspicuous, rhizines monomorphic P.flavescens
4. Maculae effigurate, rhizines dimorphic P. neotropicam
5 (3) Thallus corticicolous, sorediate but not forming pustules, medulla C-, with
hypoconstictic acid, without gyrophoric acid P. mirandum
5 Thallus saxicolous, with soredia-producing pustules, medulla C+ rose, with gyrophoric
acid, without hypoconstictic acid 6
6 (5) Thallus lobate, lobes 1.0-4.5 mm wide, cilia frequent or totally absent in some lobes
P. nylanderi
6 Thallus laciniate, laciniae . 8-3 .0 mm wide, abundantly ciliate P. pectinatum
7 (2) Upper surface with effigurate maculae, rhizines dimorphic P. subcaperatum
7 Upper surface without maculae, rhizines monomorphic 8
8 (7) Thallus corticicolous, cilia up to 0.7 mm long, medulla C-, without gyrophoric acid
P. masonii
8 Thallus saxicolous, cilia up to 2.8 mm long, medulla C+ rose or C-, gyrophoric acid
present in variable concentration 9
9 (8) Thallus lobate, lobes wide (5-15 mm), apothecial disc usually lacerate, amphithecium
rugose to scrobiculate P. magnum
9 Thallus sublaciniate, sublaciniae narro w (1.0-3.5 mm), apothecial disc entire,
amphithecium smooth P. delicatulum
10 (1) Medulla yellow P. enteroxanthum
1 Medulla white, sometimes faint ro se (decompo sition of the salazinic acid) 11
11 (10) Medulla UV+ yellow (lichexanthone), cespitose cilia abundant 12
1 1 Medulla U V- (without lichexanthone); cespito se cilia uncommon 15
12 (1 1) Thallus without vegetative propagules, usually with apothecia P. lichexanthoniciim
12 Thallus with isidia or soredia, with or without apothecia 13
13 (12) Thallus isidiate P. ultralucens
13 Thallus sorediate 14
14 (13) Soredia produced in botryose structures P. spinibarbe
14 Soredia not produced in botryose structures P. pontagrossense
15(11)
15
16(15)
16
17(15)
17
18(17)
18
19(18)
19
20(19)
20
21(19)
21
22(21)
22
23 (22)
23
24 (22)
24
25(18)
25
26 (25)
26
27 (26)
27
28 (27)
28
29 (28)
29
30 (27)
30
31 (30)
31
15
Thallus with lacinules forming arbuscular structures 16
Thallus without arbuscular structures 17
Arbuscula ciliate and esorediate, upper surface with effigurate maculae, cilia abundant
P. coralliforme
Arbuscula eciliate and sorediate, upper surface with punctiform maculae, cilia sparse
P. ramusculum
Upper surface with ciliate warts P. verrucisetosum
Upper surface without ciliate warts 18
Upper surface without maculae 19
Upper surface with effigurate, reticulate or extensive (hypermaculate) maculae 25
Thallus without vegetative propagules 20
Thallus withisidiaor soredia 21
Lobes eciliate, conidia sublageniform P. latissimum
Lobes ciliate, conidia filiform P. mantiqueirense
Thallus isidiate P. concurrens
Thallus sorediate 22
Asco spores large, 25-35 umlong, 23
Asco spores médium- sized, 12-17 umlong 24
Lobes eciliate or withrare simple cilia P. cristiferum
Lobes conspicuously ciliate, cilia simple to furcated P. sieberi
Thallus with lacinules, conidia short-filiform P. maximum
Thallus without lacinules, conidia sublageniform P. stiippenm
Upper surface with maculae primarily effigurate (a few extensive may occur) 26
Upper surface with reticular or extensive maculae, never effigurate 43
Thallus without vegetative propagules 27
Thallus withisidiaor soredia 33
Undersurface brown or with a blackish-brown center 28
Undersurface jet-black 30
Cilia usually furcated, maculae strong, rhizines monomorphic P. bifiáum
Cilia rarely furcated, maculae weak to distinct, rhizines dimorphic 29
Undersurface extensively light brown, usnic acid absent P. embescens
Undersurface brown to blackish-brown, usnic acid present P. subcaperatum
Thallus with lacinules, medulla with norlobaridone P. cristobaliae
Thallus without lacinules, medulla without norlobaridone 31
Rhizines dimorphic P. reparatum
Rhizines monomorphic 32
16
32 (31) Cilia simple P . permaculatum
32 Cilia usually furcated P. expansum
33 (26) Thallus isidiate 34
33 Thallus sorediate 35
34 (3 3 ) Undersurface abundantly veined, usnic acid present, norlobaridone absent
P. neotropicum
34 Undersurface slightly rugose to papillate, usnic acid absent, norlobaridone present,
P. sabtinctorium
35 (33) Soralia growing from the lobe margins towards the undersurface 36
35 Soralia strictly marginal, or on the upper surface, then submarginal, capitate or growing
in pustuloid formations 37
36(35) Thallus lacinulate, rhizines monomorphic, conidia unciform P. parahypotropum
36 Thallus without lacinules, rhizines dimorphic, conidia filiform P. subsumptum
37 (3 5 ) Undersurface light brown to dark brown, rarely blackish-brown in central áreas 38
37 Undersurface black 40
38 (37) Cilia frequently furcated, maculae weak P. petropoliense
38 Cilia simple, maculae distinct to strong 39
39(38) Maculae strong , rhizines monomorphic , under marginal zone erhizinate
P. leucosemothetum
3 9 Maculae distinct, rhizines dimorphic, under marginal zone rhizinate P. reitzii
40 (37) Thallus laciniate, cilia simple, up to 5 mm long P. paramoreliense
40 Thallus lobate to sublaciniate, cilia simple or furcated, up to 2 mm long 41
41 (40) Cilia furcated frequent P. concors
41 Cilia exclusively simple 42
42 (41) Lacinules abundant, often with subapical soralia P. margaritatum
42 Lacinules absent, soralia laminai or marginal P. psendoreticulatum
43 (25) Thallus without isidia, soredia, pustules or schizidia 44
43 Thallus with isidia, soredia, pustules or schizidia 51
44 (43) Thallus laciniate, laciniae narrow (up to 3 mm) 45
44 Thallus lobate or wide-laciniate (2-5 mm or even wider) 46
45 (44) Cilia simple P. warmingii
45 Cilia branched common P. herrei
46 (44) Thallus with conspicuous lacinules 47
46 Thallus without lacinules or with a few sublacinules 48
47 (46) Lacinules with acute ápices, ornate with black outlined pycnidia P. eurysacum
17
47 Lacinules often spatulate or with the apex enlarged, pycnidia without black outline
P. austromaculatum
Cilia usually squarrose P. radiatum
Cilia simple or rarely furcated 49
Under marginal zone naked, apothecia imperforate P. ruptum
Under marginal zone rhizinate, apothecia perforate 50
Maculae reticular, conidia 10-14 um P. cetratum
Maculae reticular to extensive (hypermaculate), conidia 6.5-9.0 um .... P. acanthifolium
Thallus with abundant flaking off areolae (schizidia) P. austrocetratum
Thallus with isidia, soredia or pustules, rarely with flaking off areolae 52
Isidia present 53
Isidia absent 54
Isidia simple to coralloid, not forming soredia P. subisidiosnm
Isidia soredioid, formed by sorediose clusters P. lacte um
Thallus with pustules, whichcan originate soredia ornot 55
Thallus without pustules, with soredia 58
Thallus with lacinules P. granulare
Thallus without lacinules 56
Cilia absent or rare P. ruminatum
Cilia frequentto abundant 57
Pustules ± capitate, forming dense agglomerations P. elixii
Pustules rugose, nor forming agglomerations P. niidum
Soredia usually formed at the ápices of lacinules, rarely submarginal 59
Soredia mostly submarginal, only a few formed at the ápices of sublacinules 60
Upper surface hypermaculate, medulla with lobaric acid P. albinatum
Upper surface with reticular maculae, medulla without lobaric acid P. clavuliferum
Cilia simple to furcated, up to 3.5 mm long, maculae strong P. livido-tesselatum
Cilia simple, rarely furcated, up to 1.5 mm long, maculae distinct 61
Under marginal zone rhizinate, conidia 8.5-12 um P. reticulatum
Under marginal zone naked or papillate, conidia 11-26 um 62
Cilia simple, up to 1.5 mm long, conidia 11-16 um P. reterimulosum
Cilia simple to rarely furcated, up to 0.5 mm long, conidia 15-26 um P . foliolosum
48
48
(46)
49
49
(48)
50
50
(49)
51
51
(43)
52
52
(51)
53
53
(52)
54
54
(52)
55
55
(54)
56
56
(55)
57
57
(56)
58
58
(54)
59
59
(58)
60
60
(58)
61
61
(60)
62
62
(61)
Parmotrema acanthifolium (Pers.) Spielmann & Marcelli, comb. nov.
Parmelia acanthifolia Pers., in Gaudichaud-Beaupré, Voyage autour du monde, execute sur les
corvettes 1'Uranie et la Physcienne, sur les ordres du Cap. L. de Freycinet, en 1817-1820,
Botanique, p. 197. 1826. - Type: Brazil, [Rio de Janeiro State, Municipality of] Rio de Janeiro,
corticicolous, leg. Gaiidichand 13 (holotype: PC!)
(Fig. 01)
Holotype description
Thallus brownish (herbarium), lobate, corticicolous, 9 cm broad. Lobes irregularly branched,
laterally overlapped, 2-7 mm wide, surface continuous, smooth, lustrous, becoming irregularly
cracked towards the center; apical zone rounded; margin crenate to sinuous, slightly undulated.
Maculae reticular to extensive (hypermaculate), sometimes inconspicuous, laminai, not
originating cracks. Cilia black, simple or rarely with lateral branchlets, up to 1 mm long, 0.02-
0.06 mm wide, abundant. Lacinules, pustules, soredia and isidia absent. Medulla white.
Undersarface impossible to describe in detail since the thallus is totally glued to a paper sheet,
apparently black, lustrous, smooth to rugose or papillate; marginal zone brown, lustrous, 1-
2 mm wide, rhizinate to papillate, with an attenuated limit; rhizines black, simple to squarrose,
up to 2 mm long, 0.01-0.05 mm wide, apparently abundant.
Apothecia cupuliform, 2-10 mm in diameter, stipitate, submarginal, margin crenate to
sinuous, amphithecium maculate, smooth or irregularly pitted, disc dark brown, epruinose,
perforate atmaturity; ascospores ellipsoid, 13.5-15 x 8.5-10 um, episporium ca 1 um. Pycnidia
submarginal, conspicuous, with or without prominent margin, abundant, ostiole black; conidia
short-filiform, 6.5-9.0 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major) and consalazinic
acid (minor).
Remarks: Parmotrema acanthifolium is distinct by the lobate thallus with reticular maculae or
hypermaculate, the short conidia (6.5-9.0 um) and the absence of lacinules.
This species is close to P. cetratum, and was already considered a synonym (Hale & Fletcher
1990). However, the hypermaculate thallus and shorter conidia (10-14 um in P. cetratum
according to Hale & Fletcher 1990) seems to be sound charaters to recognizeP. acanthifolium as
a good species.
Parmotrema ruptum, another close species, has imperforate apothecia, longer conidia (10-
12.5 um), shorter ascospores (10-12.5 x 6.0-7.5) and a distinct bare marginal zone on the
undersurface.
Apparently one of the first authors to accept Parmelia acanthifolia as a good taxon was
Vainio (1890), applying this name to specimens gathered by him in Sitio (nowadays
Municipality of António Carlos, Minas Gerais State). Afterwards, Mílller Argoviensis (1893)
reported the species to Costa Rica, Hue (1895) to Califórnia (U.S. A.), Zahlbruckner (1909)
included it in his key to the Brazilian Parmelia, and Lynge (1 924) mention it for Uruguay.
The record of Vainio (1890) was questioned by Hale (1960), who stated that it was based on
Parmelia subcaperata Kremp. About Lynge (1924), Hale (1960) stated that the species was
probably destroyed in Berlin during the Second World War.
In the protologue (Gaudichaud-Beaupré 1826) only one specimen was cited, and so it is the
holotype (Vienna Code, article 9.1 Note 1), not a lectotype as suggested by Hale & Fletcher
(1990).
19
Distribution: U.S.A (Hue 1895), Costa Rica (Míiller Argoviensis 1893), Uruguay (Lynge 1924)
and Brazil, where it was recorded to the States of Minas Gerais (Vainio 1890) and Rio de Janeiro
(Gaudichaud-Beaupré 1826).
Parmotrema albinatum (Moon, Kurok. & Kashiw.) Blanco, Crespo, Divakar, Elix &
Lumbsch
Mycologia97(l): 157.2005.
Rimelia albinata K.H. Moon, Kurok. & Kashiw., Journal of Japanese Botany 76 (6): 322. 2001.
- Type: U.S.A., Hawaii, Molokai Island, Molokai District, below Puu Kolekole, 21°05' N,
156°54' W, on soil, elevation about 1150 m, November 4, 1997, leg. Y. Ohmura 4210 (holotype:
TNS!;isotype:BISH).
(Figs. 02-03)
Holotype description
Thallus whitish, sublaciniate, loosely adnate, terricolous, 6.5-10 cm broad; sublaciniae
irregularly branched, crowded, 2-5 mm wide; surface continuous to minutely notched, rarely
cracked, smooth to irregular, lustrous, pruinose in the lobes ápices; apical zone rounded; margin
crenate to sublacinulate, undulated. Lacimdes simple to furcated, localized in the whole thallus,
usually plane, sometimes slightly canaliculated, apex acute when without soredia, although
frequently sorediate, then capitate or growing backwards, 0.5-2.5x0.3-1.5 mm, underside
beige. Maculae "absent". Cilia black, simple, thin, 0.5-1.2 x 0.02-0.05 mm, ± abundant. Soralia
concolor to the thallus, capitate, extensive or labriform, formed on the lacinules ápices or in
some margins, which become involute; soredia subgranular. Isidia and pustules absent. Medulla
white. Undersarface black (beige under the lacinules and sorediate lobes), lustrous, papillate,
rarely with scars; marginal zone dark brown, lustrous, 1.5-2.0 mm wide, rhizinate or sometimes
without rhizines, with an attenuated limit, smooth to papillate; rhizines black, simple, branched
near the ápices or squarrose, 0.5-2.0 x 0.01-0.06 mm, abundant, ± evenly distributed. Apothecia
absent. Pycnidia laminai to submarginal, especially abundant in the lacinules, conspicuous,
frequent, ostiole black; conidia filiform, 7.5-10.0 * ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin, salazinic and trace of lobaric acid (Moon et ai. 2001).
Remarks: Parmotrema albinatum is characterized by the sorediate and short-lacinulate thallus
with medullar salazinic acid and traces of lobaric acid. The upper surface is distinctly whitish,
probably caused by the very thick upper córtex.
It is cio se to P. clavuliferum, a species producing long lacinules and which instead of lobaric
acid presents only fatty acids (Kurokawa 1979).
We described maculae "absent" since the upper córtex is as whiter as the maculae normally
are in Rim elia-like species, and what one can see inP. albinatum is just an inconspicuous surface
relief in a reticular pattern.
The specimen preserved in TNS is labeled as the holotype, and one of its parts was illustrated
in the protologue, so probably the isotype is in BISH. This information is contrary to that
published in the same protologue (Moon et ai. 2001). There is no statement if the three parts in
the holotype collection are from the same thallus, and only one of them was illustrated in the
20
protologue. They are, notwithstanding, similar in every respect and seem to be really parts of the
same specimen. A label with the holotype, made by K.H. Moon in 1998, gives the following
chemistry: atranorin, norlobaridone and salazinic acid. One other label (undated but with
Kurokawa's handwriting), asserts that the specimen Ohmura 4210 has atranorin, salazinic acid
and lobaric acid (++). The last information is taken, by the moment, as the correct since it was
published in the protologue.
The terricolous habit of the holotype (Fig. 03) is very rare in this group of species (apparently
just known in P. herreí). However, in the protologue one saxicolous and one corticicolous
specimens are additionally listed. So maybe the substrate, in this case, is not an important feature
to be considered.
Distribution: Hawaiian Islands (Moon et ai. 2001).
Parmotrema austrocetratum Elix & Johnston
Mycotaxon 31 (2): 495. 1988.
Type: New Zealand, North Island, Taranaki Land District, New Plymouth, Burgess Park, 60 m,
on tree trunk in remnant forest, 07.V.1980, leg. J.A. Elix 7979 (holotype: CHR!; isotype:
CANB!).
= Rimelia aastrocetrata (Elix & Johnston) Hale & Fletcher, The Bryologist 93 (1): 26. 1990.
(Figs. 04-05)
Holotype description
Thallus beige (herbarium), lobate, loosely adnate, corticicolous, 12.5 x 8.0 cm broad, lobes
irregularly branched, laterally overlapped, 6-9 mm wide; surface lustrous, pruinose in some
places, reticulate or cracked, the reticulations giving rise to a pattern resembling a mosaic of
areolae, the areolae can develop revolute margins and finally ílake off (schizidia), showing the
medulla, in the central part of the thallus convex areolae can be seen, usually associated with a
strong rugosity of the thallus; apical zone rounded; marginal zone undulate, margin crenate.
Maculae distinct, reticular, laminai, originating areolae mosaics. Cilia black, simple or
sometimes squarrose, 0.1-1.6 x 0.02-0.05 mm, abundant in some lobes. Lacinules simple or
little branched, plane to canaliculated, apex usually acute, sometimes rounded or truncate, 0.5-
7.0 x 0.2-1.4 mm. Pustulae, soredia and isidia absent. Medulla white. Undersurface not seen
(thallus pasted in a paper card).
Apothecia cupuliform to urceolate, lacerate when well developed, 2.0-14.0 mm in diameter,
stipitate, laminai to submarginal, margin smooth to slightly crenate, sometimes denticulate,
amphithecium maculate, smooth to scrobiculate, disc pale brown, epruinose, perforate only in
some very old apothecia; ascospores ellipsoid, 12.5-15.0 x 8.5-10.0 um, episporium 1.0-
1.5 um. Pycnidia submarginal to laminai, especially abundant in some lacinules, conspicuous,
usually without prominent margin, ostiole black; conidia filiform, 8.0-25.0 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin, chloro atranorin, salazinic acid, consalazinic acid and trace of protocetraric
acid (Elix & Johnston 1 988).
21
Remarks: Parmotrema anstrocetratum is distinguished by the formation schizidia (Fig. 05) and
lacinules, and the presence of salazinic acid.
Elix & Johnston (1988) described the process of wrinkles formation by means of pressured
areolae of the upper córtex, which also can flake out, and called them dactyls. However, the
more appropriate term is schizidia, as already suggested by Moon et ai. (2001), who placed P.
austrocetratum as a synonym of P. hawaiiense, a position we do not agree, since P. hawaiiense
do not develops lacinules, has smooth upper surface and smaller conidia (10-15 um).
Another cio se species is the eciliate and pustulate P. ruminatum (see comments under this
species).
The holotype of P. austrocetratum is a well developed specimen in excellent state of
preservation. Notwithstanding, is totally glued by the undersurface to a paper card, so that it's
impossible to describe the underside characters. Near the specimen there's a label with the
chemical composition: atranorin, chloroatranorin, salazinic acid (major) and consalazinic acid,
determined by TLC, the analyst being PM. A. This information differs a little from the published
one, where traces of protocetraric acid were also noted.
The isotype (CANB) is also glued by the underside. Unfortunately, as no additional
specimens of P. austrocetratum were seen, the undersurface features could not be studied.
In correct number of the holotype is J.A. Elix 7979, not J.A. Elix 4645, as published in the
protologue (Elix & Johnston 1988). This aspect was also annotated by Streimann (label with the
isotype).
It was very hard to find pycnidia with conidia. After some attempts, very short ones were
found (7-12,5 um). Yet, on a final artempt very long ones (up to 25 um long) were found. As far
as we know, this is a new record in Parmotrema, followed by Parmotrema arteagum (conidia up
to 23 um, according to Egan 1982).
Distribution: Austrália and New Zealand (Elix & Johnston 1988, Galloway 2007).
Parmotrema austromaculatum Spielmann & Marcelli, sp. nov.
Type: Brazil, Rio Grande do Sul State, Municipality of Boqueirão do Leão, Linha Sinimbuzinho,
Perau da Nega, 29°20'17.2"S, 52°26'33.6"W, 430 m, saxicolous, left margin of the stream, close
to the vertical slope, open place, 23.11.2004, leg. A.A. Spielmann & L.S. Canêz 1111 (holotype:
SP).
(Fig. 06)
Holotype description
Thallus greenish gray, lobate, loosely adnate, saxicolous, 18 cm broad. Lobes irregularly
branched, laterally overlapped to crowded, 5-30 mm wide, surface smooth to reticulately
cracked, sometimes pruinose, opaque to sublustrous, becoming notched and reticulately cracked
towards the centre; apical zone rounded, plane to strongly concave; margin crenate to broadly
crenate, ascendant or revolute, undulated; lateral margin usually ascendant and revolute, regular
to undulated, often lacinulate. Maculae distinct, reticular or hypermaculate, laminai, originating
cracks or not. Cilia black, usually simple, rarely furcated, sinuous, often ascendant, especially at
the lobe ápices, 1.0-3.5 x 0.03-0.10 (-0.15) mm, abundant. Lacinules plane to concave, simple
to furcated or irregularly branched, sometimes subpalmate, often spatulate or with the apex
enlarged, rounded or concave, or apex acute to truncate, abundant, 0.5-6.0 x 0.5-7.0 mm, lower
surface black, brown, or rarely white variegated. Pustules, soredia and isidia absent. Medulla
white. Undersurface black, lustrous, mainly smooth to sometimes rugulose or slightly papillate,
rarely veined, with cracks; marginal zone dark brown, occasionally black or rarely pale brown or
22
white variegated, lustrous, 1-7 mm wide, naked, with an attenuated limit, smooth to rugulose or
papillate; rhizines black, usually simple, sometimes furcated or irregularly branched, 0.3-4.0 x
0.02-0.20 mm, frequent, distributed in groups.
Apothecia concave to cupuliform or ± urceolate, 2-6 mm in diameter, stipitate, submarginal,
margin smooth to crenulated or incised, amphithecium maculate, smooth to rugose, disc brown,
epruinose, entire to lacerated, perforate at maturity; ase osp ores broadly ellipsoid, 10.0-12.5 x
8.0-9.0 um, episporium ca. 1.0 um. Pycnidia submarginal, conspicuous, with prominent margin,
frequent especially in the lacinules, ostiole black; conidia filiform, (7.5-) 9.0-1 1 .5 x ca. 1 .0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C: atranorin, salazinic acid and consalazinic acid.
Remarks: P armo trema aiistromaculatum is characterized by the distinct reticular maculae, often
hypermaculate, conspicuous lacinules, wide lobes (up to 30 mm) and long cilia (up to 3.5 mm).
Similar to Parmotrema enrysacum (Hue) Hale, which can be differentiated by the lacinules
with submarginal pycnidia ornate with a black outline.
The species is abundant at the type locality. Additional specimens studied revealed only
minor differences: thalli varied from 6 to 30 cm broad. Lacinules were 40.4-20.0 x 0.5-8.0 mm.
Cilia reached up to 5 mm of length and conidia were 7.5-14.0 um long.
Distribution: known only from the type locality.
Paratypes: Brazil, Rio Grande do Sul State, Municipality of Boqueirão do Leão, Linha
Sinimbuzinho, Perau da Nega, 29°20'02.2"S, 52°26'27.9"W, 440 m alt, on rock at stream
margin, open place, 24.11.2004, leg. A.A. Spielmann & L.S. Canêz 698, 705 (SP); the same,
29°20'17.2"S, 52°26'33.6"W, 430 m ali, saxicolous, left margin of the stream, near the steep
slope, open place, 23.11.2004, leg. A.A. Spielmann & L.S. Canêz 1340, 1341, 1342, 1343, 1344,
1345, 1358, 1365 (SP).
Parmotrema bifidum Spielmann & Marcelli, sp. nov.
Type: Brazil, Mato Grosso State, Serra da Chapada [nowadays Chapada dos Guimarães], Bocca
da Serra, ad rupes , 15.VI.1894, leg. G.O.A. Malme (holotype: S!).
(Figs. 07-10)
Holotype description
Thallus milky grayish, lobate, loosely adnate, saxicolous, 11 cm broad. Lobes irregularly
branched, laterally overlapped, 3-7 mm wide, with a distinct black line, thickened at the sinuses,
surface continuous, smooth, lustrous, becoming strongly cracked and sometimes rugose towards
the center, the cracks usually perpendicular to the lobe axis; apical zone rounded; margin
sinuous-incised, undulated. Maculae strong, effigurate, laminai, originating cracks. Cilia black,
simple to usually furcated, sometimes more branched, up to 1 mm long, usually thick, 0.05-
0.15 mm wide, abundant. Lacinules, pustules, soredia and isidia absent. Medulla white.
Undersurface pale brown, opaque to sublustrous, usually smooth, slightly rugose or papillate in
some áreas, without cracks; marginal zone dark brown, shiny, 1-3 mm wide, naked, with an
attenuated limit, smooth or rarely papillate; rhizines dark brown, simple or less frequently
23
furcated or irregularly branched, up to 2 mm long, usually thickened, (0.2-) 0.5-0.20 mm wide,
abundant, irregularly distributed, leaving several áreas nude.
Apothecia cupuliform, 1.5-9.0 mm in diameter, substipitate, submarginal, margin usually
involute, smooth to irregularly incised, amphithecium maculate, rugose-foveolate, disc pale to
dark brown, epruinose, perforate or imperforate; ascospores ellipsoid, 11-14 x 6.5-7.5 um,
episporium 0.5-1.0 um. Pycnidia submarginal or sometimes advancing over the lamina,
conspicuous, without prominent margin, abundant, ostiole black; conidia short-filiform, 6-10 x
ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), protocetraric acid (trace).
Remarks: Parmotrema bifidum is distinguished by the thickened furcated cilia, strong effigurate
maculae, and the brown underside with a naked and shiny dark brown marginal zone. It must be
noted that the under marginal zone is darker than the center, a rare pattern inside Parmotrema
with salazinic acid (present also inP. petropoliense).
Parmotrema expansum also develops furcated cilia and effigurate maculae; however, has a
distinct shiny jet-black undersurface.
The shiny under marginal zone and branched cilia can be compared withP. lichexanthonicum,
which contain lichexanthone and has weak maculae. Strongly effigurate maculae occur in
P. leucosemothetum, distinguished by the simple longer cilia (up to 3 mm).
The cilia of P. bifidum resemble those of Parmotrema consors (Nyl.) Krog & Swinscow, a
common Brazilian species that contains only fatty acids (Spielmann & Marcelli 2009). On the
other hand, in this species the cilia are usually turned down, while in P. bifidum they are
prostrate or ascending.
Lynge (1914), when describing Parmelia cetrata Ach. subsp. radiata Lynge, listed two
specimens, one from Paraguay and another from Brazil. Later on, Hale (1960) made the
typification of this taxon choosing the specimen from Paraguay as lectotype. Hale & Fletcher
(1990) placed P. cetrata subsp. radiata as synonymous of Parmotrema cetratum. This taxon
actually constitutes a good species, P. radiatum (see comments under it). The Brazilian syntype
in not conspecific with it, and it is here being described asP. bifidum.
Distribution: Known only from the type locality.
Parmotrema cetratum (Ach.) Hale
Phytologia28 (4): 335. 1974.
Parmelia cetrata Ach., Synopsis Methodica Lichenum: 198. 1814. - Type: U.S.A.,
Pennsylvania, leg. Muhlenberg s.n. (lectotype: H-ACH!; duplicate of the lectotype: UPS,
photo!).
= Rimelia cetrata (Ach.) Hale & Fletcher, The Bryologist 93 (1): 26. 1990.
(Fig.ll)
24
Lectotype description
Thallus brownish (herbarium), laciniate, corticicolous (fide Acharius 1814), 3.5 cm broad.
Laciniae irregularly branched, 2-5 mm wide, surface continuous, smooth, lustrous, becoming
reticulately cracked towards the center; apical zone slightly rounded to acute; margin smooth,
plain to slightly undulated. Maculae distinct, reticular, laminai, originating cracks. Cilia black,
simple to furcated, up to 1.3 mm long, 0.02-0.05 (-0.10) mm wide, abundant. Lacinules,
pustules, soredia and isidia absent. Mediãla white. Undersurface black, lustrous, smooth, with
cracks; marginal zone absent; rhizines black, apparently simple or squarrose, up to 1 mm long,
abundant.
Apo the chim plane (probably pressed), 1.7 mm in diameter, probably stipitate and laminai,
margin incise to deeply incised, disc blackish brown, epruinose, perforate; ascospores ellipsoid,
12.5-15 x 8-10 um, episporium 1.0-1.5 um. Pycnidia laminai, conspicuous, without prominent
margin, frequent, ostiole black; conidia not found (10 pycnidia examined).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema cetratum is recognized by the reticular maculae, the absence of
propagules, the presence of salazinic acid, and absence of lacinules.
This species is part of a complex and misunderstood group, and several good, but frequently
abundantly lacinulate species has been synonymized under P. cetratum based on the combination
of a small set of characters usually observed.
While studying the rich lichenized mycota of Southern Brazil, Fleig (1997) and Eliasaro
(2001) found significant differences in the maculation of the specimens. Spielmann (2005)
asserted that some morphologically distinct specimens could be good taxa. Divakar et ai. (2005)
through molecular data, also pointed out the necessity to revise P. cetratum and its synonyms,
since specimens from Costa Rica tentatively identified as this species were morphologically and
molecularly distinct from the Uruguayan ones.
The revision of the types of the synonyms showed that significant morphological differences
were hidden under the name P. cetratum, a species that probably is much more restricted in its
distribution, maybe centered in Eastern North America and perhaps extending to other temperate
climates. Therefore, we are proposing some accepted synonyms (see Hale & Fletcher 1990) as
good species: Parmelia acanthifolia Pers., Parmelia ce trata Ach. subsp. radiata Lynge,
Parmelia herrei Zahlbr and Parmelia warmingii Vainio. See the key and comments under each
of the se taxa for more detailed information.
Other synonyms are placed as nomina inquirenda, since the types were not seen or, chiefly
because their size or preservation condition, do not have sufficient well-preserved characters to
clarify their status: Parmelia cinerascens var. saxicola Rãs., P. maura Pers., P. menziesii C.W.
Dodge, P. perforata var. microspora Rãs, P. perforata var. replicata Meyen & Flotow and
P. polita Fr.
The lectotype of P. cetratum (H) is a poor specimen 3.5 cm broad, although with a well-
developed apothecium, mixed with one Parmelia sp. and glued in a paper. It was consequently
very difficult to study the undersurface. Moreover, the interpretation of thallus morphology was
difficult: we called the thallus laciniate, although it could be well only part of a larger lobed
specimen, since signs of another glued specimen are visible.
Unfortunately, conidia were not found in the lectotype. Future studies including the duplicate
of the lectotype (UPS) and additional specimens recently discovered in the PH Herbarium
25
(Lendemer & Hewitt 2002) are fundamental to verify this feature, since Hale & Fletcher (1990)
did not inform the specimen from which the ascospores and conidial data were furnished.
The herbarium UPS sent a photograph of the duplicate of the lectotype of P. cetrata,
represented by a quite larger thallus, apparently without apothecia, but with more developed
laciniae. It seems to agree well with the lectotype. The same Parmelia sp. is also growing with
the specimen so it is probably from the same locality or maybe it was before glued together with
the lectotype.
The typification of P. cetratum was made by Hale & Fletcher (1990). We do not know,
however, how these authors concluded that the type locality was Pennsylvania, since both the
protologue and the lectotype (H) indicate only "America Septentrionalis".
Because the proposed changes, at the present we cannot have a real idea of the "true"
distribution ofParmotrema cetratum. The references below are maintained, however, as a guide
to the current literarure on this name.
Distribution (probably unreal): Oceania (Elix 1994, Galloway 1985, Malcolm & Galloway
1997, Louwhoff & Elix 1999), Ásia (Awasthi 1976, Kurokawa & Lai 2001), Africa (Krog &
Swinscow 1981, Swinscow & Krog 1988, Hale & Fletcher 1990, Aptroot 1991), North America
(Hale & Fletcher 1990, Brodo et ai 2001, Nash & Elix 2002c, Esslinger 2008), Central America
(Feuerer 2008) and South America (Hale & Fletcher 1990). In South America it is known to
Argentina (Lynge 1924, Osório 1976, Hale & Fletcher 1990, Adler 1992, Calvelo & Liberatore
2002), Bolivia (Feuerer et ai 1998), Brazil (Marcelli 2004), Chile (Feuerer 2008), Paraguay
(Lynge 1914), Peru (Feuerer 2008), Uruguay (Lynge 1924; Osório 1972, 1992b, 1995, 2003) and
Venezuela (Vareschi 1973). In Brazil it was recorded to the States of Minas Gerais (Lynge 1914,
1924; Hale & Fletcher 1990, Ribeiro 1998), Mato Grosso (Lynge 1914), Paraná (Osório 1977a/b,
Fleig 1997, Eliasaro 2001), Rio de Janeiro (Lynge 1924), Rio Grande do Sul (Spielmann 2006),
Santa Catarina (Marcelli 1992, Osório 1997, Fleig 1997) and São Paulo (Marcelli 1991, 1992;
Ribeiro 1998).
Parmotrema clavuliferum (Rãs.) Streimann
BibliothecaLichenologica22: 93. 1986.
Parmelia clavulifera Rásánen, Annales Botanici Societatis Zoologicae Botanicae Fennicae 20
(3): 4. 1944. Type: Oceania, Tahiti, ad corticem arboris, 1868, leg. Dr. med. Bouffon vel E.
Vieillard (lectotype: H!; paratype: H!).
= Rimelia clavulifera (Rãs.) Kurok., Journal of Japanese Botany 66 (3): 158. 1991.
(Fig. 12)
Lectotype description
Thallus brownish (herbarium), lobate, apparently loosely adnate (pasted in a paper card),
corticicolous, 12.5 cm broad. Lobes irregularly branched, more or less crowded, 2.0-7.5 mm
wide, surface continuous to cracked, smooth to irregularly pitted (i.e., with small depressions),
lustrous, becoming strongly cracked in the center; apical zone rounded; margin crenate, plane to
undulated. Lacinules simple to dichotomous, localized mainly in the center of the thallus, usually
canaliculated and "closing" (i.e., joining the margins and becoming solid) from the ápices to the
bases, apex acute and with a black tip when esorediate, capitate to extensive and finally "T-like"
when sorediate, 0.5^1.0 (-7.5) x 0.2-1.0 mm; undersurface cream color, erhizinate or with
sparse rhizinae. Maculae distinct, reticular, laminai, originating cracks. Cilia black, simple or
rarely subsquarrose, 0.2-1.5 x 0.02-0.04 mm, few. Soralia capitate to sometimes growing
26
backwards, on the lacinules ápices; soredia subgranular to farino se. Isidia and pustules absent.
Mediãla white. Undersurface (difficult to analyze, since the thallus is pasted in a paper card)
black, lustrous, slightly rugose to papillate; marginal zone brown to dark brown, lustrous, 2.5-
3.0 mm wide, naked to more usually rhizinate, with an attenuated limit, papillate; rhizines black,
simple or sometimes squarrose, up to 3 mm long, ca. 0.05 mm wide, abundant, distributed in
groups.
Apothecia absent. Pycnidia laminai, mainly on the lacinules, conspicuous, frequent, ostiole
black; conidia filiform, 7.5-12.5 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema clavuliferiim is recognized by the reticular maculae and the abundant
closed lacinules with the soralia at the ápices.
Some authors already realized that it is different from P. reticulatum (e.g. Streimann 1986,
Kurokawa 1991, Moon et ai 2001, Elix 2001, Killmann & Fischer 2005), while others (e.g.
Awasthi 1976, Divakar et ai. 2001) reached the conclusion that it is conspecific with P.
reticulatum. Based on the types here studied, we can only conclude they are two distinct species.
The lectotype (H) is well-preserved and developed. The cilia are very few. However, many
rhizinae protrude from the lower surface, and look like cilia. The thallus is pasted on a paper
card, so the description of the undersurface was not complete. Although a small portion of one
thallus, also numbered "2" like the major one, is pasted by the upper surface, it is difficult to
assign it P arme lia clavulifera on the absence of lacinules. So we preferred to take the data just
from the bigger thallus.
The correct information from the types are not clear. Rásánen (1944) asserted that the
collections were from Tahiti, made by Dr. med. Bouffon or E. Vieillard. Hale & Fletcher (1990)
also listed Tahiti, but just Vieillard as the collector. In the lectotype packed there is an annotation
by T. Ahti, dated February 2002, indicating that the collector is probably E. Vieillard, and the
locality is possibly New Caledónia.
The paratype (H) is represented by a more laciniate, robust thallus (the lectotype is more
delicate), with rhizines strongly squarrose-branched. The cilia also are more distinctly squarrose
than that of the lectotype. The lacinules are similar, however, and clearly assigns this species
within the circumscription oíParmelia clavulifera.
Several records from Brazil (e.g. Spielmann 2005, Benatti 2005 and Jungbluth 2006) were
based on different lichens that were not yet studied in detail. Actually they can represent a
different, although widely distributed species (see comments under P. cetrata f sorediifera in
the Nomina inquirenda).
Distribution: Oceania (Lamb 1963, Feuerer 2008), Ásia (Feuerer 2008), Hawaii (Moon et ai.
2001) and Brazil (Barros & Xavier Filho 1972).
Parmotrema concors (Kremp.) Spielmann & Marcelli, comb. nov.
Parmelia concors Kremp., Verhandlungen der zoologisch-botanischen Gesellschaft in Wien 30:
329-342. 1881 . -Type: Austrália, Kings Island, on bark, leg. Neate s.n. (holotype: M!).
(Fig. 13)
27
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, corticicolous, 9.0 x 7.0 cm. Lobes
irregularly branched, crowded, (2.5-) 4.0-7.0 mm wide, surface continuous or subscrobiculate,
sometimes cracked, smooth, lustrous, becoming strongly cracked in the center; apical zone
rounded; margin crenate to sublacinulate, undulated. Lacinules hardly distinct, often simple or
bifurcate, sometimes spatulate, localized mainly in the thallus center, usually plane, apex
rounded or acute, 1.0-2.5 x 0.5-2.0 mm. Maculae distinct, effigurate, laminai, forming dense
groups and sometimes assuming an reticular look in the lobe ápices. Cilia black, often stout and
spiky, straight of curved, simple to furcated, 0.2-1.5 x 0.02-0.1 mm, abundant. P ustules and
isidia absent. Soralia usually brown, labriform to growing backwards, usually subapical and
submarginal turning lobes margins involute, or in the lacinules ápices, or sometimes orbicular
and laminai; soredia granular. Medulla pale salmon (maybe herbarium color - possibly white
when fresh). Undersurface black, lustrous, rugose, papillate, with cracks; marginal zone dark
brown, lustrous, 1.0-3.0 mm wide, naked, with a attenuated limit, smooth to rugose or finely
papillate; rhizines black or sometimes with brown ápices, simple or rarely branched, slender or
thickened, 0.2-2.0 x 0.01-0.15 mm, abundant, almost evenly distributed.
Apothecia unknown. Pycnidia not found, although numerous black, submarginal spots are
present, yet without conidia.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema concors can be distinguished by the effigurate maculae, the naked under
margin and the spiky, often furcated cilia.
This species was treated as a synonym under P. reticulatum (Hale & Fletcher 1990).
Nevertheless, the effigurate maculae, the thickened, often bifurcate cilia, and the dimorphic
rhizines, leave no doubt that it's a typical Canomaculina in the Elix (1997) sense. It has also a
distinct naked brown marginal zone. Parmotrema reticulatum has slender cilia, sometimes with a
small lateral branch, but not furcated at ali, and a rhizinate marginal zone, with a very narrow
brown naked margin.
Known distribution: Austrália (Krempelhuber 1881, Hale & Fletcher 1990).
Parmotrema concurrens Hale
Mycotaxon5(2):432. 1977.
Type: Brazil, Pernambuco State, Brique, May 1970, saxicolous, leg. L. Xavier Filho 752
(holotype: US!).
(Fig. 14)
Holotype description
Thallus grayish, lobate, loosely adnate, saxicolous, pieces up to 5 cm broad. Lobes irregularly
branched, laterally overlapped to ± crowded, 3-10 mm wide, surface continuous, smooth,
sublustrous to opaque, sometimes becoming irregularly cracked in the center; apical zone
rounded; margin crenate to sublacinulate, undulated. Maculae absent. Cilia black, simple, up to
28
1 mm long, 0.02-0.05 mm wide, few. Lacinules, pustules and soredia absent. Isidia concolor to
the thallus, apex dark brown to black, usually simple, sometimes branched or aggregated in small
coralloid formations, up to 0.3 mm high, 0.05-0.20 mm wide, erect, firm to friable, apex rarely
ciliate, laminai to marginal. Medulla white. Undersurface black, opaque, usually finely rugulose,
less frequently smooth or papillate, with cracks; marginal zone brown or rarely pale brown,
sublustrous, \-4 mm wide, naked, with an attenuated limit, smooth, papillate or slightly rugose;
rhizines black, simple or rarely furcated or irregularly branched, up to 2 mm long, 0.02-0.10 mm
wide, frequent, distributed in groups.
Apothecia absent. Pycnidia submarginal or laminai, conspicuous, with or without prominent
margin, frequent, ostiole black; conidia short-filiform, 6-10 x ca. 1 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (minor), salazinic acid (major), consalazinic acid
(minor), gyrophoric acid (minor), hypoconstictic acid (minor), norhypoconstictic acid (minor).
Remarks: Parmotrema concurrens is characterized by the isidiate thallus without maculae and
the combination of salazinic acid, gyrophoric acid and hypoconstictic acid and allied in the
medulla.
The similar P. idtralacens can be distinguished by the presence of lichexanthone (medulla
UV+ yellow) and absence of gyrophoric acid. It also can resemble some saxicolous P. tinctorum
(Nyl.) Hale with narrow lobes, although not related at ali, since this last species is eciliate and
has lecanoric acid.
The holotype collection is formed by several thalli, or parts of the same thallus. Hale (1977)
reported in the protologue thallus 5-8 cm broad, however it is possible he included
measurements from the additional specimens examined. The medulla is slightly rose in some
áreas, perhaps an effect of the decomposition of the salazinic acid.
Distribution: Venezuela (Feuerer 2008), and Brazil, where it was reported to Pernambuco State
(Hale 1977).
Parmotrema coralliforme (Hale) Hale
Phytologia28 (4): 335. 1974.
Parmelia coralliformis Hale, Contributions from the United States National Herbarium 36 (5):
324. 1965. - Type: México, State of Chiapas, Mature pine forest on mountainside, 2340 m alt,
18 Km southeast of San Cristo bal on highway 190, on deciduous tree, 24 .III. 1960, leg. M.E.
Hale & T.R. Soderstrom 20287 (holotype: US ! ; isotypes: REN, S ! ).
(Fig. 15)
Holotype description
Thallus pale brownish (herbarium), lobate, loosely adnate, corticicolous, 12.5 cm broad. Lobes
irregularly branched, laterally overlapped to crowded, 3-12 mm wide, surface continuous,
smooth, lustrous, becoming cracked towards the center; apical zone rounded; margin broadly
crenate to incised, undulated. Maculae distinct, effigurate, laminai, originating effigurate cracks.
Cilia black, simple to sometimes furcated or rarely dichotomous, up to 1.5 mm long, sometimes
very thick, 0.04-0.15 mm, abundant. Lacinides forming arbuscular structures up to 10 mm high,
coralloid-branched, branches flattened to terete, usually ciliate, surface scrobiculate, not
29
sorediate but with abundant nodules specially in the ápices. Pustules, soredia and isidia absent.
Medulla white. Undersurface (based in the S isotype): dark brown, lustrous, smooth to papillate
or rarely with inconspicuous veins, without cracks; marginal zone yellowish brown, lustrous, 2-
12 mm wide, rhizinate, with an attenuated limit, smooth; rhizines black, simple to squarrose, up
to 1 mm long, thin, 0.01-0.05 mm wide, abundant, evenly distributed.
Apothecia absent. Pycnidia submarginal or rarely laminai, conspicuous, without prominent
margin, frequent, ostiole black; conidia not found (ca. 30 pycnidia examined).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema coralliforme is distinguished by the arbuscular structures, which are
usually ciliate and esorediate, the effigurate maculae and the pale brown and rhizinate marginal
zone in the undersurface.
It can be compared with P. ramusculum, which develops eciliate and strongly sorediate
arbuscular structures, has punctiform maculae, sparse cilia (abundant in P. coralliforme) and the
black naked marginal undersurface. See underP. ramusculum comments about other species.
In P. coralliforme, the effigurate maculae give rise to distinct cracks, also effigurate. This
phenomenon is not common in the species here investigated, since the cracks are usually
irregular or reticular, independent from the type of macula.
The arbuscular structures sometimes break, exposing the white medulla, but true soredia are
totally absent.
The isotype (S) agrees well with the ho lo type. It has more ciliate fascicular structures and, as
the holotype, abundant pycnidia; however, no conidia were found. Since the holotype is glued to
a card, the isotype was used to describe the underside.
Distribution: Guianas (Sipman 1999) and México (Hale 1965).
Parmotrema cristiferum (Taylor) Hale
Phytologia 28 (4): 335. 1974.
Parmelia cristifera Taylor, London Journal of Botany 6: 165. 1847. - Type: índia, Calcutta,
Wallich s.n. (lectotype: FH!).
= Parmelia perforata var. ulophylla Meyen & Flotow, Novorum Actorum Academiae Caesareae
Leopoldino-Carolinae Naturae Curiosum 19, suppl. 1: 218. 1843. - Type: Hawaiian Islands,
Oahu (lectotype: G!).
= Parmelia hildebrandtii Kremp., Linnaea 41: 139. 1877. - Type: Cômoro Islands, Johanna
Island, 400 m. alt, an der Rinde Von Báumen (besonders Cycas), an hehten stellen im Walde,
leg. Hildebrandt 1866c (holotype: M!; isotypes: BM!, K!).
= Parmelia mesotropa Mull. Arg. f sorediosa Míill. Arg., Flora 74: 377. 1891. - Type: índia,
Cachar, leg. Keenan (holotype: G!).
(Figs. 16-17)
30
Lectotype description
Thallus cream, lobate, apparently loosely adnate, 16 x 14 cm. Lobes strongly undulated,
irregularly branched, laterally overlapped and difficult of to distinguish one from the other, 10-
18 mm wide., surface usually cracked; apical zone rounded and crisped; margin strongly
sorediate and crispate-undulated. Maculae absent. Soralia whitish, linear continuous or
interrupted, sometimes labriform, mainly marginal, although some present in thalline crests,
rarely orbicular or capitate, or in small "columns"; soredia farinose to granular. Laciniães, cilia,
pustules and isidia absent. Medulla white or with the upper part slightly rose (maybe because the
eumitrins). Undersurface black, lustrous, strongly papillate, with scars; marginal zone absent or
cream under the soralia, opaque, 1-2 mm wide, naked, with a sharp limit, smooth or rugose;
rhizines black, simple, up to 1 mm long, few, distributed in groups.
Apothecia concave, small (up to 2.5 mm in diameter), substipitate, laminai, margin smooth,
sorediate, amphithecium sorediate, disc brown, epruinose, imperforate; ascospores ellipsoid, 24-
35 x 12-20 um, episporium 2-3 um. Pycnidia apparently rare (there are several black spots that
remember pycnidia, however without any tissue inside); conidia not found (ca. 30 pycnidia
studied).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC (J.A. Elix, 28.X.1998, label with the holotype in FH): atranorin (minor),
chloroatranorin (trace), salazinic acid (major), consalazinic acid (trace), eumitrin Al (minor),
eumitrin A3 (trace), unknowns (minor/trace).
Remarks: Parmotrema cristifemm is characterized by the upper córtex with atranorin, linear
marginal soralia, sublageniform conidia and large ascospores (24-35 x 12-20 um). It is a usually
eciliate species, although cilia can rarely be found in some specimens.
It can be compared with P. stuppeum that also develops marginal soralia and sublageniform
conidia, but otherwise has smaller ascospores (12-17 x 6-9 um, fide Hale 1965).
Parmelia cristifera was first proposed by Taylor (1847), who cited four specimens, without
designating the type. They are, therefore, syntypes: "Calcutta, Wallich; Mauritius, Dr. Wright;
Brazil, Gardner; Demerara, Mr. Parker". Krog & Swinscow (1981) already dealt with this
difficulty, and it seems appropriate to see each specimen separately:
- índia, Calcutta, Wallich s.n., was selected as the lectotype of Parmelia cristifera (see Berry
1941, Dodge 1959 and Hale 1965).
- Brazil, Gardner, is the holotype of Parmelia gardneri C.W. Dodge (Dodge 1959).
- Mauritius, Dr. Wight (not "Wright" as cited in the protologue) represents one syntype of
Parmelia cristifera. And, at the same time, is one specimen cited in the protologue of
Parmelia sieberi (Dodge 1959).
- Demerara (Guyana), Mr. Parker, is not Parmelia cristifera, since the salazinic acid is lacking
(medulla K-). There' s a label from Krog pointing out the existence of only fatty acids (TLC),
and another one from Hale (May 08, 1959), determining this species as Parmelia sanctae-
cmcis. Krog & Swinscow (1981) asserted that it belongs to Parmelia praesorediosa.
Seemingly the first author to choose the specimen Wallich s.n. (FH!) as lectotype was Berry
(1941). He left a label (signed E.C.B.) in 1938, pointing this. Nevertheless, the ascospores
measures (6-8 x 2-A um) published in 1941 are certainly a typographical error, since so small
ascospores are not known in Parmotrema.
Lindsay (1866) made comments about the Gardner specimen from Brazil, i.e., Parmotrema
gardneri (Dodge) Hale. Muller Argoviensis (1888) asserted that Calcutta specimen was sterile,
while the Brazilian one, fertile. However, the two types (FH!) are clearly fertile. Merrill (1909)
31
reported Parmelia cristifera to the United States and Hawaii, distinguishing it of P. latíssima
Fée.
Lynge (1914), when describing Malme's collections from Mato Grosso State, Brazil, found
ascospores with 22-27 x 10.5-14 um, with an episporium 2 um wide. These are little smaller
than that published by Hale (1965): 26-35 x 13-18 um, episporium 3-A um.
Lamb (1963) cited one variety and two forms to Parmelia cristifera: P. cristifera var.
abissinica Cengia Sambo is actually Flavopunctelia flaventior (Stirton) Hale (Hale 1980, Hale &
DePriest 1999). Parmelia cristifera f. pallida Rás. is a synonym of Parmotrema perlatum (Hale
1965) and finally P. cristifera f. cinerata Zahlbr. corresponds to Parmotrema cristiferum (Hale
1965).
This is one of the cases where Hale (1965) choose illustrate additional specimens examined
instead the type material, one from México (Hale 20073, Fig. 5) and the other from Formosa
(Taiwan) (Ogata s.n., Fig. 16). These were not asked on loan. Even so, we prefer to choose the
lectotype from FH to illustrate here.
The type is glued by the underside, so only some marginal zones were studied.
We failed in find conidia in the lectotype. Vainio (1909a) observed that "Secundum specimen
no. 35399 in herb. Nyl. var. cristifera apotheciis instructa est cupuliformibus, breviter
pedicellatis, imperforatis, disco testaceo rufo, sporis long. 0,026-0,032 mm., crassit. 0,015-0,018
mm, pycnoconidiis sublageniformibus, long. 0,005-0,007 mm., crass. 0,001 mm (secund. annot.
Nylanderi in specimine a Jardin lecto e loco incerto)".
It is difficult to know the substrate of the syntypes. The protologue (Taylor 1 847) gives no
indication.
Notes about accepted synonyms
Parmelia perforata var. ulophylla Meyen & Flotow
We have studied from G Herbarium two specimens. One is sorediate ("A"), sterile, with strongly
crisped lobes and occasional cilia (up to 0.5 mm long) in one part of the thallus. So it clearly
matches with Parmotrema cristiferum, when compared with the lectotype (FH!). The occasional
cilia were realized by Meyen & Flotow (1843).
The second thallus ("B") has two imperforate apothecia, reticular maculae and cracks, being a
Rimelia with salazinic acid (medulla K+ yellow — > red). There is an anonymous annotation of
the ascospores (14-17 x 8-11 um), confirming they are very different from those of P.
cristiferum. The actual status of this specimen remains unclear and was not further investigated
here, since it is in a poor condition.
Parmelia hildebrandtii Kremp.
Krempelhuber (1877) describedP. hildebrandtii with ascospores 11-15 x 11-12 um. This size is
very distant from the known in Parmotrema cristiferum. Krog & Swinscow (1981) already
wondered about the identity of this species and pointed out it could be compared with P.
stuppeum on the ascospores size basis.
Hale (1965) stated that are isotypes in BM and K, but do not indicated the location of the
holotype, and was in this mode of citation folio wed by Winnem (1975). Actually, the holotype of
P. hildebrandtii is located in M, as we disco vered after consultation of the curator, Dr. Andreas
Beck. It is a well-developed specimen, composed by three thalli, two of them certainly part of
the same thallus, and the third part with several apothecia (as indicated by Krempelhuber 1 877).
Seven attempts to find ascospores were made in the more mature apothecia, but fruitless.
Conidia, however, are sublageniform, 5-7 x ca. 1 um, so matching the conidia of P. cristiferum.
In the exsiccate Krempelhuber left a drawing of the asei with ascospores inside, only two
outside. The sizes given are 1 1-15 x 1 1-12 um.
32
In the apothecium from the K specimen the hymenium is strongly guttulate and little
developed, since only few ascospores were found, some clearly immature. They are 20-29 x 10-
14 um, the episporium 2-2.5 um, cio se to the accepted to P. cristiferum. If new collections from
the type localities become available in the future, maybe the difference in the ascospores sizes
will be still smaller or absent.
Unfortunately the pycnidia are absent in the K specimen. The specimen from BM has
submarginal pycnidia with sublageniform conidia 6.5-8.5 x ca. 1 um, agreeing to the accepted
for P armo trema cristiferum and equal to that found in the holotype of P. hildebrandtii .
In conclusion, probably Krempelhuber found immature ascospores, and therefore P arme lia
hildebrandtii is synonym of Parmotrema cristiferum.
With the BM specimen there's an annotation made by B. Winnem, dated 22.11.1974,
informing that atranorin and salazinic acid were found (TLC).
Parmelia hildebrandtii Kremp. f sorediosa Míill. Arg.
Hale (1965) referred this synonym with one lectotype in G and one isotype in K. The curator of
K Herbarium informed that some specimens were not sent because could be deposited at the
Natural History Museum.
The specimen received from G consists in a small, sorediate thallus with sublageniform
conidia, 5-8 x ca. 1 um. It is morphologically similar to P. cristiferum, and thus we agree with
the synonymization proposed by Hale (1965). However, the K specimen could provide
additional evidences.
Parmelia mesotropa Míill. Arg. f sorediosa Míill. Arg. (Fig. 17)
Hale (1965) asserted that the holotype of this taxon would be deposited in K, while the isotype in
G. We have received two specimens from these herbaria. The specimen in G is labeled as
holotype (contrary to the information given in Hale 1965). Morphologically and chemically it
agrees well with the lectotype of P. cristiferum, and so is accepted here as belonging to it. The
chemistry was reported in a label by Elix (3. X. 1998), through TLC in solvent C and HPLC:
atranorin (major), chloroatranorin (trace), salazinic acid (major), consalazinic acid (minor),
eumitrin Al (trace), eumitrin A3 (trace).
The specimen from K sent to us has marginal soredia and a naked under margin, clearly a
Parmotrema, without cilia. The córtex is K+ yellow, UV-, the medulla K-, C+ red, KC+ red, P-.
Since it has lecanoric acid, probably belongs to Parmotrema austrosinense (Zahlbr.) Hale. The
conidia are filiform, 9-13 x ca. lum. There's no annotation from Hale, so maybe he don't saw
this specimen. There's also no indication if this is a type or not.
Therefore, by the moment we are accepting the G specimen as the holotype. It is possible that
the real type was not sent and is preserved in K Herbarium. This needs to be further checked.
Distribution: Africa (Hue 1899, Hale 1965, Krog & Swinscow 1981), Ásia (Taylor 1847,
Vainio 1909a, Hue 1899, Hale 1965, Chen et ai 2005, Divakar & Upreti 2005), Oceania (Merrill
1909, Hue 1899, Hale 1965, Elix 1994, Louwhoff & Elix 1998), North America (Merrill 1909,
Berry 1941, Hale 1965, Brodo et ai 2001), Central America (Hue 1899, Hale 1965) and South
America (Hue 1899, Hale 1965). In South America it is known to Brazil, British Guiana,
Colômbia, Peru, Surinam and Venezuela (Hale 1965). In Brazil it was recorded to the States of
Minas Gerais (Zahlbruckner 1905), Mato Grosso (Lynge 1914), Minas Gerais (Zahlbruckner
1905), Paraná (Donha 2005) and São Paulo (Zahlbruckner 1909, Hale 1965).
33
Parmotrema crisíobaliae (Ferraro & Elix) Blanco, Crespo, Divakar, Elix & Lumbsch
Mycologia97(l): 157.2005.
Rimeliella crisíobaliae Ferraro & Elix, Mycotaxon 49: 406. 1993. - Type: Argentina, Comentes
Province, Depto. Ituzaingó, Rincón Ombú Chico, leg. L.I. Ferraro, S. Tressen, A. Schinini & M.
Gonzales 687, 25. IV. 1975 (holotype: CTES!).
= Canomacidina crisíobaliae (L.I. Ferraro & Elix) Elix, Mycoíaxon 65: 477. 1997.
(Fig. 18)
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, corticicolous, 10.5 cm broad. Lobes
irregularly branched, ± crowded, 3-10 mm wide, surface continuous to cracked, smooth to
notched, lustrous, becoming reticulate cracked towards the center; apical zone rounded to
abundantly lacinulate; margin deeply crenate to sublacinulate, undulated. Maculae distinct,
effigurate, although sometimes assuming a pseudo-reticular aspect, laminai, originating cracks or
not. Cilia black, simple to rarely squarrose or furcated, usually short, up to 0.7 mm long, 0.02-
0.04 mm wide, abundant. Lacinules irregularly branched to subdichotomous, present in the
whole thallus, plane to slightly convex, apex rounded, truncate or acute, 2-7 x 1-6 mm. Isidia,
soredia and pusíides absent. Medulla white. Undersurface black, lustrous, smooth or papillate,
with cracks; marginal zone brown to dark brown, lustrous, 1-3 mm wide, rhizinate and papillate,
with an attenuated limit; rhizines black, dimorphous, short rhizines simple, up to 0.5 mm long,
0.01-0.04 mm wide, abundant, evenly distribute, long rhizines usually subsquarrose to
irregularly branched, up to 2 mm long, 0.04-0.06 mm wide, frequent, distributed in groups.
Apoíhecia ± plane, 6-10 mm in diameter, stipitate, margin smooth to slightly incise,
undulated, amphithecium maculate, rugose, disc brown, epruinose, perforate; ascospores
ellipsoid, 12-14 x 7.5-9.0 um, episporium 1.0-1.5 um. Pycnidia laminai or more commonly
submarginal, conspicuous, with or without prominent margin, abundant, ostiole black; conidia
filiform, 9-14 x ca. 1.0 um.
Color íesís: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvení C, HPLC: atranorin, chloroatranorin, salazinic acid (major), norlobaridone
(major), consalazinic acid (minor/trace), loxodin (minor/trace) (Ferraro & Elix 1993).
Remarks: Parmoírema crisíobaliae is distinguished by the strongly lacinulate thallus without
vegetative propagules, the effigurate maculae, dimorphic rhizines and the production of salazinic
acid and norlobaridone.
Together the effigurate maculae, this species also presents pseudo-reticular ones. Since it has
a black undersurface (common in Rimeliá), not brown as usual in Rimeliella, it could well link
between the se formerly recognized genera.
Ribeiro (1998) reported P. crisíobaliae to MG, Brazil. However, the study of the specimen
Marcelli & Yano 23972 (SP) shows that it belongs to another taxon, probably P. subcaperaíum.
In this way, P. crisíobaliae remains known only to Argentina and Bolivia.
The holotype from CTES is a good specimen, fertile and fully lacinulate. The apothecia are
variably damaged, giving the idea of an old thallus. Also some features of the lobes are difficult
to evaluate since the thallus is, for the mo st part, covered by the lacinules.
The correct orthography of the specific epithet is "crisíobaliae" instead of "crisíobalif\ as
pointed out by Ribeiro (1998) and Hale & DePriest (1999). It follows the Article 60.11 and
Recommendation 60C.1 of the Vienna Code.
34
Distribution: in South America it was recorded to Argentina (Ferraro & Elix 1993) and Bolivia
(Ferraro& Elix 1993).
Parmotrema delicatulum (Vainio) Hale
Phytologia28 (4): 335. 1974.
Parmelia delicatula Vainio, Acta Societatis pro Fauna et Flora Fennica 7(1): 35. - Type: Brazil,
Minas Gerais State, [Municipality of] Catas Altas, Caraça [nowadays Parque Natural do Caraça],
1885, leg. Vainio (holotype: TUR-V! isotypes: BM!, FH!, M, UPS).
= Parmelia radians Lynge, Arkiv for Botanik 13 (13): 85. 1914. - Type: Brazil, Minas Gerais
State, Municipality of São João d'el Rey [dei Rei], leg. G.O.A. Malme 203 (holotype: S!;
isotype: MO).
(Fig. 19)
Holotype description
Thallus yellowish, blackening toward the centre, sublaciniate, loosely adnate, saxicolous, 4.5-
5.0 cm broad. Sublaciniae irregularly branched, contiguous with laterally overlapped or crowded
margins, 1.0-3.5 mm wide, surface continuous, smooth, lustrous, becoming cracked in the
center; apical zone rounded, or subtruncate; margin crenate to sublacinulate, undulated. Cilia
black, simple or sometimes furcated, 1.5-2.8 x 0.03-0.10 mm, abundant in the young regions.
Lacinules, maculae, pustules, soralia and isidia absent. Medulla white. Undersurface black,
lustrous, smooth, papillate or slightly rugose, with numerous scars; marginal zone usually
absent, i.e., black, or rarely dark brown, lustrous, 1.0-1.2 mm wide, naked, smooth or with few
papillae; rhizines black, simple or rarely furcated next to the apex, 0.2-1.5 x 0.01-0.05 mm,
frequent, irregularly distributed.
Apothecia cupuliform, 1.3-4.3 mm in diameter, substipitate, submarginal, margin smooth or
slightly crenate with small incisions, amphithecium smooth, pruinose, maculate, eciliate, disc
dark brown, epruinose, imperforate; ascospores ellipsoid, 11-12.5 x 7.5-9.0 fim, episporium
0.5-1.0 um. Pycnidia submarginal, rarely laminai, conspicuous, with or without prominent
margin, abundant, ostiole black; conidia short-filiform, straight or slightly curved, 7.5-9.0 x ca.
0.5 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC (J.A. Elix, 07.V.1996, label with the holotype in TUR-V): usnic acid
(minor), salazinic acid (major), consalazinic acid (minor), gyrophoric acid (minor),
hypoconstictic acid (minor), lecanoric acid (trace).
Remarks: Parmotrema delicatulum can be recognized by the yellowish thallus (usnic acid)
composed of narrow sublaciniae (1.0-3.5 mm wide) and developing long cilia. The closest
species is P. magnum, distinguished by the wider lobes (5-15 mm), shorter cilia (up to 1.5 mm)
and the abundance of medullar gyrophoric acid (C+ rose). Additionally, the amphithecia are
smooth inP. delicatulum and strongly rugose inP. magnum.
The holotype of P. delicatulum has the upper surface blackening towards the centre, but it
seems derived from the herbarium preservation or the age of the specimen.
35
The BM isotype agrees well with the features of the holotype, but only immature ascospores
were found. The isotype from M was not found by the curator, and could be on loan (A. Beck,
pers. comm.). According to Hale (1965), there's an isotype of P arme lia delicatula in UPS,
however, this specimen was not sent to us, and only a picture could be seen.
The holotype of Parmelia radians has a strongly pruinose upper surface, cilia more
commonly furcated and a more plane and adnate thallus than Parmotrema delicatulum. His
chemical study (TLC in solvent C, HPLC - J.A. Elix, 10.V.1996) revealed the presence of usnic
acid (minor), salazinic acid (major), consalazinic acid (minor) and hypoconstictic acid (trace).
This profile makes it also more similar to P. delicatulum than to P. magnum, which contains
gyrophoric acid (major) and does not produces hypoconstictic acid. Its conidia are short-filiform,
6-9 x 1 um, matching both P. delicatulum and P. magnum. Unfortunately, apothecia are not
known in P. radians, so that no additional features could be compared and we are accepting this
taxon as synonym of P. delicatulum, by the moment.
As far as we know, this is the first publication illustrating the type of Parmelia delicatula.
Distribution: South America, where is reported to Argentina (Osório 1981), Brazil
(Zahlbruckner 1909, 1930a; Hale 1965; Fleig 1997), Paraguay (Osório 1970), Uruguay (Hale,
1965) and Venezuela (Feuerer 2008). In Brazil it is known to the States of Mato Grosso do Sul
(Fleig & Riquelme 1991), Minas Gerais, Paraná, Rio de Janeiro (Hale, 1965) and Rio Grande do
Sul (Spielmann 2006).
Parmotrema elixii Spielmann & Marcelli, nom. nov.
Type: Austrália, Western Austrália, Mt Clarence, Albany, 35°02'S. 117°54'W, 230 m ali, dry
sclerophyll forest with large granite outcrops, on shaded granite ledges, 15.IX.1994, leg. J.A.
Elix,H.T. Lumbsch & H. Streimann 41377 (holotype: CANB; isotype: CANB!, HO, PERTH!).
= Rime lia pustulata Elix & Bawingan, Mycotaxon 81: 252. 2002.
= Parmotrema pustulatum (Elix & Bawingan) O. Blanco, A. Crespo, Divakar, Elix & Lumbsch,
Mycologia 97 (1): 157, 2005, non Parmotrema pustulatum Louwhoff & Elix, Mycotaxon 75:
199. 2000.
(Fig. 20)
Isotype description (CANB)
Thallus brownish (herbarium), lo bate, loosely adnate, saxicolous, 6-9 cm broad. Lobes
irregularly branched, ± crowded, 2-10 mm wide, surface continuous or cracked, smooth or
slightly rugose, lustrous, becoming rugose and reticulate cracked towards the center, in some
áreas the areolae ílaking off; apical zone rounded; margin smooth to crenate or sublacinulate,
undulated. Maculae distinct, reticular, laminai, originating cracks. Cilia black, simple, short, up
to 0.5 mm long, 0.02-0.05 mm wide, frequent. Lobules spatulate, frequently formed from the
pustules, 0.5-2.0 x 0.4-1.5 mm. P ustules ± capitate, usually forming dense agglomerations,
laminai and submarginal, originating soredia-like structures. Lacinules, isidia and true soredia
absent. Medulla white. Undersurface black, lustrous, usually papillate or rugose, with cracks;
marginal zone dark brown, lustrous, 1-3 mm wide, rhizinate, with an attenuated limit, papillate;
rhizines black, simple to branched, up to 1.5 mm long, 0.02-0.10 mm wide, frequent, irregularly
distributed.
Apothecia unknown. Pycnidia immature, submarginal, inconspicuous, without prominent
margin, rare, ostiole black; conidia not found (10 pycnidia examined).
36
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC (J.A. Elix, 07.XI.2002, label with the isotype in CANB): atranorin
(minor), chloroatranorin (minor), salazinic acid (major), consalazinic acid (minor).
Remarks: Parmotrema elixii is distinguished by the reticular maculae, the medulla with
salazinic acid, short cilia (up to 0.5 mm long) and principally by the production of pustules,
usually forming dense agglomerations.
It can be compared with P. granulare that has also reticular maculae but otherwise distinct
lacinules with individualized pustules (i.e., not forming agglomerations), and P. anchietanam
Benatti, Marcelli & Elix (see Benatti et ai 2008), characterized by the narrow (1-3 mm) and
lacinulate lobes
The new name P. elixii is here being proposed to the species earlier known as Rimelia
pustulata Elix & Bawingan (Elix et ai 2002). That species is clearly different from Parmotrema
pustulatum Louwhoff & Elix (Louwhoff & Elix 2000), which is emaculate and without salazinic
acid. When Blanco et ai (2005) synonymized the genus Rimelia under Parmotrema, they
proposed the combination "Parmotrema pustulatum (Elix & Bawingan) O. Blanco, A. Crespo,
Divakar, Elix & Lumbsch" to Rimelia pustulata, so creating a later homonym and, therefore, the
necessity of a new name.
We did not have access to the holotype, however CANB sent an isotype. It is well developed
and ali features could be described. In the isotype from PERTH, however, the underside was not
studied, since the lichen is glued to a paper card. Even so, it agrees well with the CANB isotype.
Distribution: Austrália (Elix et ai 2002), Philippines (Elix et ai 2002).
Parmotrema enteroxanthum Hale
Mycotaxon 5 (2): 434.1977.
Type: Venezuela, Merida, La Carbonera, sector El Pedregal, on rocks in open pasture, 2100 m
Alt., 19-111-1975, leg. M.E. Hale & M. Lopez Figueiras 44177 (holotype: US; isotypes: MERF,
UPS!).
(Fig.21)
Isotype description (UPS)
Thallus whitish-grey, lobate, loosely adnate, saxicolous, 9 cm broad; lobes irregularly branched,
crowded, 0.5-1.3 mm wide, surface continuous to irregularly cracked, sometimes flaking off
some parts of the upper córtex, smooth, lustrous to opaque, becoming cracked in the center;
apical zone rounded; margin smooth or irregular, undulated. Maculae and cilia absent. True
isidia absent. Dactyloid structures concolor to the thallus, simple to coralloid, up to 1 mm high,
usually thick and inílated, 0.2-0.8 mm wide, erect, firm or friable, then leaving a yellow tuft of
medulla, apex eciliate Pustules laminai, extensive, giving raise to schizidioid or dactyloid
structures. Lacinules and soredia absent. Medulla entirely yellow or more frequently only the
under portion. Undersurface black, lustrous, smooth, slightly rugose or papillate, rarely veined,
strongly cracked; marginal zone absent (black) or pale to dark brown, lustrous, 1.5-4.0 mm
wide, naked, with an attenuated to sharp limit, smooth; rhizines black, simple or rarely branched,
up to 1.5 mm long, 0.01-0.10 mm wide, frequent, distributed in groups.
Apothecia cupuliform, 1.0-7.5 mm in diameter, stipitate, laminai to submarginal, margin
crenate to incised, usually involute, amphithecium smooth to strongly pustulate, emaculate or
37
with faint maculae, disc brown, epruinose, imperforate; ascospores ellipsoid to slightly reniform
30-35 x 14-19 um, episporium 2.5-4.5 um. Pycnidia mainly submarginal, conspicuous, without
prominent margin, abundant, ostiole black; conidia sublageniform, 5.0-6.5 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor)
Remarks: Parmotrema enteroxanthum can be recognized by the yellow medulla (at least the
inferior portion), swollen and dactyloid isidia, sublageniform conidia and large ascospores
(30-35 um long).
The holotype collections from US was not sent to us, the same with the isotype from MERF.
The isotype in UPS is composed of two parts which, presumably, are part of one single thallus.
But it is interesting to note that one part has well developed isidia, while the other not. In the
other fearures, however, the two parts are identical, including the yellow medulla and large
ascospores.
As far as we know, there are no other species of Parmotrema with salazinic acid and yellow
medulla.
Distribution: Colômbia (Aguirre-C & Rangel-CH 2007), Venezuela (Hale 1977).
Parmotrema erubescens (Stirton) Krog & Swinscow
TheLichenologistl5(2): 127-130. 1983.
Parmelia erubescens Stirton, Scottish Naturalist 4: 201. 1878. - Type: Queensland, near
Brisbane, leg. F.M. Bailey 1 1 (lectotype: BM!, selected here).
= Canomaciãina erubescens (Stirton) Elix, Mycotaxon 65: 477. 1997.
(Fig. 22)
Holotype description
Thallus brown reddish (herbarium), lo bate, loosely adnate, 3-A cm broad. Lobes irregularly
branched, laterally overlapped, 2-10 mm wide, surface continuous, smooth to scrobiculate,
lustrous, becoming rugose, cracked and more deeply scrobiculate towards the center; apical zone
rounded; margin broadly crenate to incised, undulated. Maculae distinct, effigurate, sometimes
the irregular dots connecting in pseudo -reticular formations, laminai, originating cracks or not.
Cilia black, simple or rarely furcated, up to 2 mm long, thin to thickened 0.02-0.06 mm wide,
abundant. Lacinules, pustules, soredia and isidia absent. Medulla white. Undersurface light
brown or blackish brown only in the very centre, lustrous, smooth to papillate, without cracks;
marginal zone absent; rhizines black, dimorphic, the shorter ones simple to furcated or less
frequently squarrose, up to 0.5 (-1.0) mm long, thin, 0.01-0.05 mm wide, abundant, evenly
distributed; the longer ones up to 1.5 mm long, thickened, 0.05-0.10 mm wide, simple or
branched, apparently frequent and distributed in groups.
Apothecia absent (according to Elix 1997, ascospores 10-12 x 8-10 um). Pycnidia
submarginal to laminai, conspicuous, without prominent margin, frequent, ostiole black; conidia
filiform, 8-1 1 x ca. 1 .0 um.
38
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema embescens can be recognized by the brown reddish distinctly
scrobiculate upper surface, effigurate maculae, dimorphic rhizines, brown undersurface and
absence of propagules.
Hale (1965) first regarded Parmelia embescens as conspecific with P. subcaperata. In this
way he was followed by Winnem (1975), Kurokawa (1991) and Elix (1994). Krog & Swinscow
(1981), however, segregated P. embescens based on the absence of usnic acid (present in P.
subcaperatum) and the presence of norlobaridone, which they found only in the African
specimens. Actually, the lichen they named as P. embescens could be another species, since the
considerations constant in the item "Circumscription of the species" and on the Table 13 (Krog
& Swinscow 1981: 156) lead to the conclusion that the specimens with salazinic acid and
norlobaridone are "unnamed". Some years later, Ferraro & Elix (2000) described Parmotrema
cristobaliae with base on this chemical combination. So the material studied by Krog &
Swinscow (1981) must be revised. These authors also referred the species to Brazil although
without citing the specimens examined.
Finally, Elix (1997) became convinced that P. embescens and P. subcaperata are distinct
species, based on several aspects (see Table and comments underP. subcaperatum).
Unfortunately, the holotype of P. embescens lacks apothecia, and the specimens seen by Elix
(1997) were not studied here. Nevertheless, the conidial size we found is very cio se to that found
by him (10-14 um) and his description also matches with the features we observed in the
holotype.
The holotype is constituted by five thalli fragments glued in a paper card. Two of them, the
larger ones, undoubtedly belonging to P. embescens, were marked "A" in the label and are here
being selected as lectotype. The three other thalli (B) belong to another species, with the upper
surface grayish green, inconspicuous maculae, few cilia, the undersurface black and brown only
in the narrow marginal zone, bacillar or slightly sublageniform conidia 5-6 x ca. 1 um, and
probably with stictic acid (medulla K+ yellow, C-, KC-, P+ orange).
The description of the underside is very incomplete, since the thalli are glued to the paper.
Distribution: Africa (Krog & Swinscow 1981), Oceania (Elix 1997) and in South America it
was recorded to Brazil (Krog & Swinscow 1981), although without a voucher.
Elix (1997), Krog & Swinscow (1981), Swinscow & Krog (1988), Winnem (1975).
Parmotrema eurysacum (Hue) Hale
Phytologia28 (4): 336. 1974.
Parmelia eurysaca Hue, Nouvelles Archives du Muséum d'Histoire Naturelle de Paris, sér. 4 (1):
194. 1899. - Type: México, 1865-1866, leg. M Borgeau (holotype: P!).
(Fig. 23)
Holotype description
Thallus brownish, lobate, loosely adnate, corticicolous, 13.5 cm broad. Lobes irregularly
branched, laterally overlapped to ± crowded in the center, 5-13 mm wide, surface continuous,
39
smooth, opaque to sublustrous, pruinose in some lobe ápices; apical zone rounded; margin
sinuous, undulated. Maculae extensive (hypermaculate). Cilia black, simple, up to 3.5 mm long,
0.05-0.10 mm wide, frequent. Lacinules simple to furcated, plane or rarely subcanaliculated,
apex acute, 3-10 x 0.4-2.5 mm, undersurface black to dark brown or sometimes with brown
stains. P ustules, soredia and isidia absent. Medulla white or slightly salmon (salazinic acid?).
Undersurface black, lustrous (satin-like), smooth or sometimes slightly rugulose, with cracks;
marginal zone brown to pale brown, lustrous, 2-5 mm wide, naked, with attenuated limit,
smooth; rhizines black or dark brown near the marginal zone, simple to richly squarrose or
irregularly branched, up to 2 mm long, 0.01-0.15 mm wide, frequent, evenly distributed,
although leaving several naked áreas.
Apothecia cupuliform, 4-22 mm in diameter, substipitate, margin involute, smooth to
irregularly incised, amphithecium maculate, smooth or minutely scrobiculate, disc brown to dark
brown, epruinose, widely perforate; ascospores ellipsoid, 12-15 x 7.5-9.0 um, episporium ca.
1 um. Pycnidia restricted to the lacinules, [sub] marginal or subapical, conspicuous, with
prominent margin and a distinct black outline, frequent, ostiole brown; conidia short-filiform, 6-
9 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), gyrophoric acid (minor).
Remarks: P armo trema eurysacum is distinguished by the conspicuous lacinules ornate with
almost marginal black outlined pycnidia, the hypermaculate upper surface, long simple cilia, and
the black undersurface with a bare, brown marginal zone.
The wide lobes apparently emaculate (hypermaculate) make this species comparable with P.
latissimum, which lacks lacinules, has larger ascospores (27-35 um long) and sublageniform
conidia. Hale (1965) compared P eurysacum with P. stuppeum, sorediate and also with
sublageniform conidia. The types of P. stuppeum studied here, however, are almost eciliate.
One interesting feature of P. eurysacum is the black distinct outline around the pycnidia,
which has, contrary to ali species here seen, brown ostiole. These pycnidia are apparently
marginal, however the ostiole open at the upper surface of the lacinules.
Fleig (1997) proposed P. ruptum andP. expansum as synonyms ifP. eurysacum. Parmotrema
ruptum has reticulate maculae (sometimes hypermaculate) and rare, usually simple and shorter
cilia (up to 0.5 mm), while P. expansum (see below) presents effigurate maculae, furcated cilia
and larger ascospores (16-20 .um).
Hale & DePriest (1999) proposed P. permaculatum as synonym of P. eurysacum.
Nevertheless, P. permaculatum lacks lacinules, has effigurate maculae, shorter cilia (up to 2
mm), and sometimes a white mottled marginal zone on the undersurface.
Based on the literature, we proposed a key and table assuming that ali species inside the
'Parmotrema eurysacum complex" are good (Spielmann 2005), which proved to be effective
after the types have been studied here.
Kurokawa (2001) asserted that in P. eurysacum the lacinules are white in the undersurface,
but this not happens with the holotype, where they are clearly black or at best dark brown or with
some brown stains.
Distribution: Africa (Krog & Swinscow 1981), Oceania (Elix 1994), North America (Hale
1965, Kurokawa 2001) and South America, where it was reported for the Brazilian States of
Minas Gerais (Kalb 1982, Kurokawa 2001) and Rio Grande do Sul (Fleig 1997).
40
Parmotrema expansum Hale
Mycotaxon5(2):435. 1977.
Type: Costa Rica, Province of San José, Patarrá, Desamparados, elevation 1300 m, 04-VII-1948,
on rocks, 1300 m alt, 04.VII.1948, leg. M Guirós Calvo 1462 (holotype: US!; isotype: UPS).
(Fig. 24)
Holotype description
Thallus stramineous, lobate, loosely adnate, saxicolous, 21 cm broad. Lobes irregularly
branched, laterally overlapped, 4-10 mm wide, with a thin to absent black line, surface
continuous, smooth to slightly irregular, lustrous, usually pruinose at the lobe ápices, becoming
cracked and sometimes rugose towards the center; apical zone rounded; margin broadly crenate
to deeply incised, undulated. Maculae distinct to strong, effigurate, sometimes very dense and
becoming hypermaculate, laminai, originating cracks. Cilia black, simple to furcated or
sometimes cespitose, up to 1.5 mm long, 0.04-0.10 mm wide, abundant. Lacinules, pustules,
soredia and isidia absent. Mediãla white. Undersurface black, shiny, smooth to slightly rugose
or reticulately veined, with cracks; marginal zone brown or stained with dark brown or black,
lustrous to shiny, 2-4 mm wide, naked, with attenuated limit, smooth, slightly rugose or
papillate, sometimes veined; rhizines black, simple or rarely branched near the apex, up to
1.5 mm long, 0.01-0.05 mm wide, abundant, evenly distributed.
Apothecia ± plane to cupuliform, 1-7 mm in diameter, substipitate, laminai, margin smooth to
dentate, lacerate on old apothecia, sometimes forming ciliate lobes, amphithecium maculate,
smooth to irregular, disc brown, epruinose, imperforate; ascospores ellipsoid, 16-20 x 9.0-
11.5 um, episporium 1.0-1.5 um. Pycnidia submarginal to laminai, conspicuous, with or without
prominent margin, abundant, ostiole black; conidia filiform, 7-12 x ca. 1 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema expansum is recognized by the effigurate maculae, commonly furcated
cilia, black undersurface and ascospores médium- sized 16-20 um long.
It can be compared with P. bifidum, which also has furcated cilia, but presents somewhat
darker and more ascendant lobes, a distinct black line in the lobe margins and the undersurface is
pale brown.
Other species with usually branched (cespitose) cilia also have lichexanthone in the medulla,
like P. lichexanthoniciim (without vegetative propagules) and P. spinibarbe (sorediate).
Hale & DePriest (1999) proposed P. expansum as synonym of P. ruptiim. However,
P. ruptum is a clearly distinct species that has reticulate maculae (sometimes hypermaculate),
rare, usually simple and shorter cilia (up to 0.5 mm) and shorter ascospores (10-12.5 um).
The specimen Marcelli 2243, identified asP. expansum by Benatti (2005) was checked. It has
similar cilia, but the overall morphology and color of the thallus, the shiny dark brown
undersurface, the absence of effigurate maculae and the shorter conidia (6-9 um according to
Benatti 2005) makes it not identifiable as P. expansum. Actually, it resembles those Parmotrema
with both salazinic acid and lichexanthone, cited above. However, TLC analysis demonstrated
only atranorin, salazinic and consalazinic acid. This chemical profile, combined with the features
above can represent a new taxon. Unfortunately, it is sterile, so ascospores data could not be
compared.
41
The holotype is constituted by two parts of the same thallus. The larger part was clearly cut at
the middle, and perhaps that piece was sent to UPS, where the isotype is located (Hale 1977).
Distribution: Costa Rica and Brazil, where it was recorded to the Minas Gerais State (Hale
1977).
Additional specimens examined: Brazil, São Paulo State, Municipality of São Sebastião,
Centro de Biologia Marinha da USP (CEBIMAR), over rocks, 19.111.1988, leg. M.P. Marcelli
2243 (SP).
Parmotrema flavescens (Kremp.) Hale
Phytologia28 (4): 336. 1974.
Parmelia glaberrima v ar. flavescens Kremp., Flora 52 (14): 223. 1869. - Type: Brazil, Rio de
Janeiro State, Municipality of Rio de Janeiro, leg. Glaziou 1833 (holotype: M!; isotype: H-NYL
35454!)
= Parmelia glaberrima f flavescens (Kremp.) Kremp., Flora 59: 73. 1876.
= Parmelia latíssima f flavescens (Kremp.) Mull. Arg., Linnaea 43: 32. 1880.
= Parmelia flavescens (Kremp.) Nyl., Flora 68: 607. 1885.
= Parmelia protoflavescens Zahlbr., Denskschriften der Mathematsch-Naturwissenschaftlichen
Klasse der Kaiserlichen Academie der Wissenschaften 83: 176. 1909. Illegitimate name based
on Parmelia glaberrima P flavescens Kremp., fide Hale (1965).
= Parmelia pseudo flavescens Zahlbr., Denskschriften der Mathematsch-Naturwissenschaftlichen
Klasse der Kaiserlichen Academie der Wissenschaften 83: 176. 1909. Nomen nudum, fide
Hale (1965).
= Parmelia mauriensis Hue, Nouvelles Archives du Muséum d'Histoire Naturelle de Paris, ser 4
(1): 201. 1899. Type: México, San Luis de Potosí, Abrededores, leg. Maury 7651 (holotype:
P!).
(Fig. 25)
Holotype description
Thallus yellowish tan (herbarium), lobate, loosely adnate, probably saxicolous, 15 cm broad.
Lobes irregularly branched, crowded, 2-10 mm wide, surface continuous, usually smooth and
lustrous; apical zone rounded; margin crenate to broadly crenate, or isidiate, undulated. Maculae
inconspicuous, but extensive, do not forming cracks. Cilia black, simple or rarely furcated at the
tip, 0.3-2.0 mm, rare. Isidia with base concolor to the córtex, apex usually brown or sometimes
concolor, simple to variously branched or coralloid, sometimes flattened, 0.1-2.0 x 0.05-0.2
mm, erect, firm, apex eciliate, mainly submarginal and marginal. Lacinules, pustules and soredia
absent. Medulla white. Undersurface black, lustrous, smooth, slightly rugose or papillate;
marginal zone brown to dark brown, lustrous, 1-5 mm wide, naked, with an attenuated limit,
smooth to slightly rugose or papillate; rhizines black, simple or furcated near the ápices, up to 3
mm long, 0.05-0.10 mm wide, frequent, distributed in groups.
Apothecia absent. Pycnidia submarginal, conspicuous, without prominent margin, few, ostiole
black; conidia short-filiform, 6-9 x ca. 1 .0 um.
42
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: usnic acid (minor), atranorin (trace), salazinic acid (major), consalazinic acid
(minor), gyrophoric acid (minor).
Remarks: Parmotrema flavescens is characterized by the presence of usnic acid, salazinic acid
and isidia, together with the presence of only monomorphic rhizines. This combination is, as far
as we know, unique in Parmotrema. Only P. neotropicum also produce isidia and has usnic acid,
but is differentiated by the effigurate maculae and dimorphic rhizines.
Related species with usnic and salazinic acids are P. delicatulum and P. masonii (without
propagules) and the sorediate P. mirandum, P. nylanderi andP. pectinatum.
The holotype of P. flavescens (M) is a well developed thallus, although somewhat friable, and
is formed by some pieces that probably were part of the same thallus. The upper surface shows a
peculiar pattern of whitish stains, which could well be characterized as maculae. However, we
are not sure about the nature of this "maculation", and maybe further anatomical studies could
bring more information on the subject.
The isotype from H is undoubtedly part of the holotype. Together there is a part of a thallus of
Cladonia, probably belonging to the C. verticillaris complex (according to Ahti & Marcelli
1995). This could indicate that the type of P. flavescens is saxicolous.
The holotype of Parmelia mauriensis is constituted by two sheets, one with only one large
thallus, and another with two thalli glued in a paper. The first was at some time already glued, as
one can see looking at the undersurface, and it has a label from Hale dated 1962, indicating it as
holotype. It is a large thallus, with more cilia and abundant isidia. Apart from this it agrees well
with P. flavescens, including the conidia type and size. The second sheet contains two thalli that
are possibly part of the same collection, since the features are very similar.
Hale (1960) placed Parmelia microdactyla in the synonymy of P. delicatula and compared it
with P. flavescens, the difference being the production of lobules instead of isidia. In fact, rare
flattened isidia can be seen in the holotype of P. flavescens. But the total absence of isidia,
together with a paler thallus can setP. microdactyla closer to P. magna.
Distribution: Africa (Míiller Argoviensis 1880), Central America (Hue 1899, Hale 1965,
Tenório et ai. 2002) and South America (Hale 1965). In South America it is known to Brazil,
where it was recorded to the States of Minas Gerais (Ribeiro 1998), Mato Grosso do Sul (Fleig
& Riquelme 1991), Paraná (Eliasaro 2001, Donha 2005), Rio de Janeiro (Krempelhuber 1869,
1876a; Nylander 1885; Hale 1965), Rio Grande do Sul (Fleig 1997) and São Paulo
(Zahlbruckner 1909, Ribeiro 1998, Benatti 2005).
Parmotrema foliolosum (C.W. Dodge) Spielmann & Marcelli, comb. nov.
Parmelia folio lo sa C.W. Dodge, Annals of the Missouri Botanical Garden 46 (1-2): 89. 1959. -
Type: Madagáscar, East Imerina, Andrangolóaka [Dodge 1959 wrote "Andrangolaoka"],
terricolous, XI. 1880, leg. J.M. Hildebrandt (Comm. C. Rensch.), (holotype: FH!).
(Figs. 26-27)
Holotype description
Thallus grayish, lobate, loosely adnate, terricolous, formed by several pieces (according with the
protologue reaching up to 6 cm). Lobes irregularly branched, laterally overlapped, 1-6 mm wide,
surface continuous, reticulate, sometimes cracked, smooth, and opaque, remaining continuous or
43
becoming rugose in the older parts; apical zone rounded, often concave; margin usually crenate,
sometimes sublacinulate, undulated. Maculae reticular, laminai, sometimes extensive
(hypermaculate), originating cracks. Cilia black, simple or rarely furcated, up to 0.5 mm long
(usually shorter), abundant. Soralia usually in shapeless extensive formations, sometimes
orbicular, often submarginal and turning the lobes involute; soredia farinose. Isidia, pustiães and
laciniães absent. Medulla white. Undersurface black, lustrous, smooth or papillate, with few
cracks; marginal zone dark brown, lustrous, 1-2 mm wide, naked or rarely rhizinate, with a
attenuated limit, smooth or papillate; rhizines black, usually simple, sometimes thickened (as in
Canomaculina) and branched, rarely squarrose, up to 1 mm long, abundant, more or less evenly
distributed.
Apothecia absent. Pycnidia usually submarginal, conspicuous, without prominent margin,
frequent, ostiole black; conidia filiform, (7-) 15-26 (-30) x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), protocetraric acid (trace).
Remarks: Parmotrema foliolosum can be recognized by the hypermaculate upper surface, short
cilia (up to 0.5 mm), long conidia (up to 30 um), black marginal zone in the upper surface, and a
brown nude marginal zone in the undersurface (Fig. 27). There's also thickened and branched
rhizines, resembling that of the former genus Canomaculina sensu Elix (1997).
This species was placed as synonym of P. reticidatum, which has only reticular maculae
(never hypermaculate) and shorter conidia (up to 12 um).
Parmotrema foliolosum has the longest conidia of the genus, followed by those of P.
austrocetratum (up to 25 um) andP. arteagum (up to 23 um, fide Egan 1982).
Distribution: Africa (Dodge 1959).
Parmotrema gt anulai e (Asahina) Spielmann & Marcelli, comb. et stat. nov.
Parmelia cetrata Ach. f granularis Asahina, The Journal of Japanese Botany 16: 593. 1940. -
Type: Formosa (Taiwan), Prov. Taichu, Keitau, 24.XII.1933, leg. Y. Asahina 33125 (holotype:
TNS!).
(Fig. 28)
Holotype description
Thallus brownish in herbarium, lobate, loosely adnate, corticicolous, 23 x 16 cm broad. Lobes
more or less radial, irregularly branched, contiguous, 4-11 mm wide, surface continuous,
smooth, lustrous, becoming rugose and reticulated cracked in the center; apical zone rounded;
margin broadly crenate, undulate or lacinulate. Laciniães canaliculated, corniculate contorted,
subdichotomous to subpalmate, localized in the whole thallus, but much more abundant in the
centre, apex usually acute, sometimes rounded, 1-12 x 0.4-2.0 mm. Maculae distinct, reticular,
laminai, originating cracks. Cilia black, simple to squarrose, (0.1-) 1.0-2.5 x 0.02-0.05 mm,
abundant. Pustules rugose to vesiculate, situated almost exclusively in the lacinules, giving rise
to soralia. Soralia pustular; soredia granular. Isidia absent. Medulla white. Undersurface black,
lustrous, papillate, with numerous scars in the centre; marginal zone brown, lustrous, 1-4 mm
44
wide, naked, with an attenuated limit, papillate or with scarce rhizines, rarely veined; rhizines
black, simple or squarrose, (0.2-) 0.5-3.0 x 0.02-0.06 mm, abundant, irregularly distributed.
Apothecia unknown. Pycnidia in the lacinules, laminai or subapical, conspicuous, with or
without prominent margin, few, ostiole black; conidia filiform, 7.5-13.0 x ca. 0.5 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin and salazinic acid.
Remarks: Parmotrema granulare is distinguished by the vesicular and rugose pustules along the
lacinules, the reticular maculae, the ciliate margin and the presence of salazinic acid. The thallus
is relatively large (23 x 16 cm), with wide lobes (4-1 1 mm).
This species was first described as Parmelia cetrata f. graniãaris (Asahina 1940).
Afterwards, Asahina (1952) placed it as a synonym of Parmelia cetrata f. subisidiosa Mull. Arg.
Apparently it was studied only by Asahina himself, and fortunately Hale & Fletcher (1990) listed
it as synonym of P. reticulatum.
However, P. reticulatum is a not pustulate species whose soralia is extensively submarginal or
situated in short lacinules (0.5-2.2 mm), has a smaller thallus 6.5-7.0 cm large, present crowded
lobes and shorter cilia (0. 1-1 .0 mm).
The holotype of P. granulare is well developed, with the necessary fearures to characterize
the species, except the apothecia. Fertile specimens, from the type locality (Taiwan) or close
locations could bring more data about this taxon, known only by the holotype.
The presence of the numerous vesicular pustules probably lead Asahina (1940) to propose the
epithet "granularis".
Distribution: Taiwan (Asahina 1940, 1952).
Parmotrema herrei (Zahlbr.) Spielmann & Marcelli, comb. nov.
Parmelia herrei Zahlbr., in Herre, Proceedings of the Washington Academy of Sciences 7: 353.
1906. - Type: U.S. A., Califórnia, Santa Cruz Peninsula, Pilarcitos Canyon, two miles from sea,
elevation 200 feet, May 28, 1904, leg. A.W.C J. Herre 516 (lectotype: FH!; duplicates of the
lectotype:FH!,W!).
(Fig. 29)
Lectotype description
Thallus beige (herbarium), laciniate, adnate, growing "on earth in the crevices of sandstone", up
to 7 cm broad; laciniae usually plane, less frequently undulate and with concavities, or
subcanaliculate, dichotomous, crowded, 1-3 mm wide; surface continuous, smooth, lustrous,
becoming cracked in the older parts; apex usually acute, sometimes truncate or slightly rounded;
margin smooth or sublacinulate. Maculae distinct, reticular, laminai. Cilia black, usually
branched (irregular, subdichotomous to subpalmate), rarely squarrose, up to 3 mm long,
thickened, contiguous. Pustules, soredia and isidia absent. Medulla white. Undersurface black,
lustrous, rugose, with a few cracks; marginal zone absent or dark brown, lustrous, up to 3 mm
wide, naked, rhizinate or papillate; rhizines black, simple or less frequently branched, up to
2 mm long, thinner than cilia, frequent, evenly distributed.
Apothecia unknown. Pycnidia laminai, more abundant in the apical zones, conspicuous,
without prominent margin, abundant, ostiole black; conidia unknown (ca. 30 pycnidia analyzed).
45
Color reactions: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (minor), salazinic acid (major), consalazinic acid
(minor), protocetraric acid (trace).
Remarks: Parmotrema herrei can be recognized by the long (up to 3 mm), thickened and
branched cilia, and the strictly laciniate thallus.
It is close to Parmotrema cetratum, and was up to know situated as its synonym (Hale &
Fletcher 1990). Nevertheless, typicalP. cetratum (lectotype: H-ACH; duplicate of the lectotype:
UPS) has a wider laciniae (2-5 mm) and shorter cilia (up to 1 .3), just simple to furcated.
Although lectotype, duplicates of the lectotype and "topotypes" are plenty of pycnidia, no
conidia could be found (numerous pycnidia examined). Maybe future collections of fresh
material could contribute with this information. Contact with North American lichenologists
(especially the Californian Lichen Society) looking for additional specimens unfortunately did
not returned results. Probably this is an endangered species, although originally mentioned as
common in the type locality (Herre 1910).
The lectotype (FH) is small although representative. It is growing with the sorediate
Parmotrema reticulatum and another Parmeliaceae. Both isotype and the "topotype" (FH) are
well developed and also identical to the holotype in every detail.
As far as we know, the only collector of P. herrei was Albert W. C. T. Herre, and this species
is known just to the type locality. Wiggins (1962) described the collection method of Herre: "On
the field trips he rode a bicycle with an old suitcase strapped to the rack and loaded his
specimens into it. He explored nearly every road, trail, canyon and mountainside on the whole
Santa Cruz Peninsular Apparently the first Herre' s collection of P. herrei was made in May 28,
1904 (lectotype in FH, duplicate of the lectotype in W). Then, again in June 30, 1906
("topotype", No. 825, FH), and in May 22, 1942 (specimen from the Herbarium Herre in F!).
The typification of this species is a little obscure. Zahlbruckner in Herre (1906) asserted that
specimens were "in the herbaria ofLeland Stanford Júnior University, Dr. A. Zahlbruckner, Dr.
H.E. Hasse, and the author...Type, No. 516". Berry (1941) stated that the type was in US,
collected in May 28, 1904, and listed also several Herre specimens made in June 30, 1906,
deposited in MBG, F, MIN, LSU and Hale & Fletcher (1990) chose one specimen deposited in
FH as the lectotype, and from W and US as "isolectotypes"
Actually, in 1963 Hale annotated "holotype" in the FH specimen, a collection that clearly
belonged to the "Herb. Hasse". Also in 1963, Hale labeled a specimen in FH (No. 516) as
isotype and the No. 825 as "topotype". The specimen in W was labeled as "holotype". There is
also a specimen in US whose packet was annotated by Hale as "topotype (compared with the
holotype)", A.C. Herre 825, June 30, 1906 [it seems to be part of the No. 825 deposited in FH].
We don't know if the Herbarium of Stanford University hold specimens of P. herrei,
especially the specimen cited in the protologue (Herre 1906). If positive, it can be selected as the
type. On the other hand, Herre (1942) asserted that the collections seen by Berry (1941) were
from his personal herbarium, not from the herbarium of Stanford University. So the typification
of this species deserves future work, and in the moment we are using, in part, the choices made
by Hale & Fletcher (1990).
The only author besides Herre to mention this species was Berry (1941). In this rarely cited
paper, P. herrei was reported as having a K- medulla. In the Field Museum (F) there is a
specimen determined by E.C. Berry, on March 4th, 1940. It is, actually, a Parmotrema with
ciliate isidia, the medulla K+ yellow (stictic acid), perhaps P. crinitum (Ach.) Choisy.
Distribution: Known only from the type locality (Herre 1906, 1910; Berry 1941).
46
Additional specimens examined: U.S. A: Califórnia, Pilarcitos Creek, Santa Cruz Mountains,
earth & sandstone, 200-300 feet, June 30, 1906, leg. A.C. Herre 825 (FH, US). Idem, Pilarcitos
Creek, Idem, Pilarcitos Creek Canon, San Mateo County, on sandstone, elev. 400-500 feet, May
22, 1942, leg. A.C. Herre (F).
Parmotrema lacteum Marcelli & Spielmann
Bibliotheca Lichenologica 96: 215. 2007.
Type: Brazil, Rio Grande do Sul State, Municipality of Boqueirão do Leão, Cascata do Gamelão,
saxicolous, on the rocky wall downstream from the cascades, inside the forest (shady),
29°18'13.7"S, 52°26'51.7"W, 500 m alt.., 01.11. 2004, leg. A.A. Spielmann Sc MA. Sulzbacher
1307(holotype: SP!).
(Fig. 30)
Holotype description
Thalhis saxicolous, milky pale gray, lobate, membranaceous, loosely adnate, 7.5-11.5 cm broad;
lobes irregularly branched, laterally overlapping, 2.5-7.0 mm wide, plane to concave, with
extended median axis and rounded ápices; margin subundulate, often ascending, crenate to
sublacinulate; lateral margin plane to ascending, lacinulate; distai surface smooth to slightly
scrobiculate, opaque, reticulate-maculate, becoming cracked in the center; lacinulae marginal,
simple, furcated or irregularly branched, 0.2-2.5 x 0.2-2.0 mm, plane to canaliculated, with a
rounded or acute apex, present over the entire thallus; maculae distinct, reticulate, laminai,
developing into cracks; cilia black, simple to commonly subsquarrose, 0.20-2.00 x 0.02-0.05
mm, frequent, evenly distributed. Pustules absent. Soredia granular, produced mostly from
subapical bulges of the lacinulae; true isidia absent; however, the soredia may pile up forming
structures (isidioid soredia) which are ecorticate, terete, not ramified, with brown ciliate apex,
0.10-0.50 x 0.02-0.10 mm (including the cilia), often marginal, sometimes submarginal and
reaching the lamina, rarely laminai, generally forming capitate sorediose clusters on the
lacinulae. Medulla mostly white, but with an orange, K - pigment at the isidia clusters. Lower
surface black, shiny, smooth to slightly rugose; marginal zone dark brown or sometimes beige,
1-2 mm wide, shiny, nude, rhizinate or papillate-rhizinate, with attenuate borders, smooth to
slightly rugose; rhizinae black, simple to squarrose or irregularly branched, 0.10-3.00 x 0.02-
0.05 mm, abundant, evenly distributed.
Apothecia and pycnidia unknown.
Color reactions: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C: atranorin, salazinic acid and consalazinic acid, unknown pigment.
Remarks: Parmotrema lacteum is recognized by the milk-white thallus coloration, the
commonly subsquarrose cilia, the capitate sorediose clusters originating isidioid soredia, the
orange pigment in the ápices of the lacinulae, and the presence of salazinic and consalazinic
acids.
Parmotrema subisidiosum (Mull. Arg.) Hale (holotype: G) is distinguished by the shorter,
simple cilia (up to 0.6 mm), and the formation of marginal to laminai, simple to coralloid true
isidia, 0.1-0.2 * 0.05-0.15 mm, that remain entire or can form granules, but not soredia.
Parmotrema bonplandii (Mata) Blanco, Crespo, Divakar, Elix & Lumbsch, produces
47
norlobaridone and lichexanthone in the medulla (Mata Garcia 1994). Finally, P. clavuliferum is
somewhat similar in forming soredia on the lacinulae, but never produces isidioid structures.
This species has now the distribution extended to São Paulo State.
Distribution: Brazil, Rio Grande do Sul State (Marcelli et ai. 2007) and São Paulo State (here
reported).
Additional specimens examined: Brazil, Rio Grande do Sul State, Municipality of Boqueirão
do Leão, Cascata do Gamelão, saxicolous on the rocky wall downstream from the cascades,
inside the forest (shady), 29°18'13.7"S 52°26'51.7"W, 500 m alt., 01.11.2004, leg. A.A. Spielmann
& M.A. Sulzbacher 748, 751 (SP). São Paulo State, Municipality of São Luís do Paraitinga,
Parque Estadual da Serra do Mar, Núcleo Santa Virgínia, near the director' s house,
23°19'43,0"S, 45°08'26,3"W, 915 m alt., corticicolous, field with forest in the surroundings,
exposed, 14.1.2007, leg. A.A. Spielmann, P. Jungbluth, L.S. Canêz & MJ. Kitaura 3362 (SP).
Idem, at Trilha da Pirapitinga, 23°20'17,2"S, 45°08'45,8"W, 915 m alt., corticicolous, forest
border, exposed, 14.1.2007, leg. A.A. Spielmann, P. Jungbluth, L.S. Canêz Sc MJ. Kitaura 3343
(SP).
Parmotrema latissimum (Fée) Hale
Phytologia28 (4): 337. 1974.
Parmelia latíssima Fée, Ess. Crypt, Suppl.: 119, pi. 38, fig. 4. 1837. - Type: Jamaica, s.c.
(lectotype: G!; duplicate of the lectotype: G!, Herbier Muller Arg.).
(Fig. 31)
Lectotype description
Thallus brownish, lobate, loosely adnate, 12.5 cm broad. Lobes irregularly branched, laterally
overlapped, very large, 6-25 mm wide, surface continuous, smooth, lustrous, becoming slightly
rugose and irregularly cracked in the center; apical zone rounded; margin smooth, undulated,
occasionally with some adventitious lobules. Maculae absent. Cilia absent. Lacinules, pustules,
soredia and isidia absent. Medulla white. Undersurface impossible to describe since the thallus
is totally glued to a card, but the large nude marginal zone is visible.
Apothecia concave to cupuliform, up to 15 mm in diameter, substipitate, laminai or
submarginal, margin incised to lacerate, often involute, amphithecium maculate, rugose to
irregularly areolate-cracked, disc dark brown, epruinose, imperforate or perforate in one
apothecium; ascospores ellipsoid, large, 27-35 x 15-17 um, episporium 2-3 um. Pycnidia
submarginal or less commonly laminai, conspicuous, with or without prominent margin,
abundant, ostiole black; conidia sublageniform, 6-8 x ca. 1 .0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (minor), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema latissimum can be recognized by the wide lobes, sublageniform conidia
and absence of cilia and propagules.
The general morphology, conidia and ascospores are similar to that of P. cristiferum. This
supports the idea they constitute a species pair, as already asserted by Hale (1965).
48
Parmotrema mantiqueirense also has wide lobes and lack propagules, but develops
conspicuous cilia and has filiform conidia.
The lectotype of P. latissimum is totally glued to a paper card, so the underside could not be
described. The typification of this species also requires some comments. From the three
specimens we received upon request, two have Hale's annotations: 1) the specimen from
"Jamaica insula" (G00053795), with the label information about his publication and which
exactly correspond to the plate published by Fée (1837), was annotated as "isotype" [actually the
lectotype] by Hale in 1962; 2) the specimen growing "ad ramos cinchonarum" (G00053796),
was annotated as "P arme lia zollingeri Hepp" by Hale in 1962.
In our opinion, the specimen G00053975 is the lectotype oí Parmelia latíssima, designated by
Hale, since it has the Hale's handwriting and exactly the same data cited by him in his
monograph (Hale 1965).
The specimen G00053796 is actually composed by three glued thalli in a card. The thallus at
the right of the card we labeled "A": it has a well developed, imperforate apothecium,
sublageniform conidia, 6-8 x ca. 1 um, ascospores ellipsoid, 27-33 x 13.0-16.5 um, episporium
2-3 um. The upper córtex contains probably atranorin (K+ yellow) and the medulla is K+
brownish, C-, KC- and P+ yellow — > quickly orange (protocetraric acid?). The thallus at the left
of the card (labeled "B") has also sublageniform conidia (6-8 x ca. 1 um) and the same reactions
as the "A" thallus. In 1962, Hale labeled this specimen as Parmelia zollingeri. However, future
studies are necessary since the list of synonyms of this species is long, and Hale (1965)
apparently employed a wide species concept, accepting both specimens ciliate and eciliate as
belonging to the same taxa (see also Elix 1998).
Finally, the thallus at the middle of the card ("C") is an isidiate lichen with the upper córtex
K+ yellow (atranorin) and the medulla K-, C+ red, KC+ red and P- (lecanoric acid), clearly
belonging to Parmotrema tinctorum or some close species.
The third specimen (G00053794) sent to us has a label from Muller Argoviensis (dated 1896)
indicating the published data from Fée (1837) and another from R.v. Aubel (dated 22.11.1985)
indicating that this is the lectotype, according to Hale (1965). But there are no Hale's annotations
with this specimen and we believe that Hale really never saw it. It is identical morphologically
(including the sublageniform conidia) and chemically with the specimen G00053975, and so this
specimen (G00053794) can be named as "duplicate of the lectotype".
Distribution: Ásia (Hale 1965, Divakar & Upreti 2005), North, Central and South America
(Hale 1965). In South America it was reported to Brazil, Colômbia, Peru, Venezuela, Hale
(1965). In Brazil it was recorded to the States of Mato Grosso (Hale 1965) and Mato Grosso do
Sul (Fleig & Riquelme 1991).
Parmotrema leucosemothetum (Hue) Hale
Phytologia 28 (4): 337.
Parmelia lencosemotheta Hue, Nouvelles Archives du Muséum d'Histoire Naturelle de Paris,
sér. 4 (1): 192. 1899. - Type: México, surroundings of San Luis Potosí, December 1886, leg. P.
Maury 7650 (holotype: P!).
= Canomacidina leucosemotheta (Hue) Elix, Mycotaxon 65: 477. 1997.
(Figs. 32-34)
49
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, 13-15 cm broad. Lobes irregularly
branched, laterally overlapped, 6-17 mm wide, surface continuous or irregularly cracked,
smooth, with some irregular depressions, lustrous, becoming rugose and reticulately cracked
towards the center; apical zone rounded; margin broadly crenate-incised or sublacinulate,
undulated. Maculae strong, effigurate, laminai, originating cracks. Cilia black, simple and
usually thickened and ± tapered, up to 2.5 (-3.0) mm long, 0.04-0.10 mm wide, abundant.
Soralia whitish, marginal or in the apex of the sublacinules, linear interrupted; soredia farinose
to subgranular. Lacinules, pustules and isidia absent. Medulla white. Undersurface dark brown
to blackish in some central áreas, shiny, smooth to rugose and scrobiculate, also with some veins,
with cracks; marginal zone absent or usually dark brown and shiny, naked; rhizines black or
concolor to the undersurface, monomorphic, up to 2 mm long, usually thickened, 0.02-0.20 mm
wide, simple or irregularly branched, especially near the ápices, abundant, distributed in groups.
Apothecia absent [according to Hale (1965), apothecia rare, ± adnate, to 6 mm in diameter,
amphithecium sorediate, disc imperforate or rarely perforate; ascospores 12-16 x 7-10 um].
Pycnidia submarginal to laminai, conspicuous, usually without prominent margin, frequent,
ostiole black; conidia filiform, 8-12 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema leucosemothetum is characterized by the soralia almost exclusively
marginal, the strongly maculate upper córtex (Fig. 33), the simple, usually thickened and ±
tapered cilia, and the distinct bare marginal zone on the undersurface (Fig. 34).
It has been constantly confused with P. subsumptum, which have the marginal soralia
growing backwards on the undersurface, scarce cilia (if present) and weak maculae.
Parmotrema petropoliense, another near species, has slender bifurcate cilia.
An interesting feature to note is that in P. leucosemothetum holotype the thickened rhizines
are much more common than the slender ones. Usually in this group (formerly Rimeliellà) the
contrary is the rale.
Krog & Swinscow (1981) reducedP. leucosemothetum to synonymy of P. subsumptum, with
base on the variation they found in African species. Fleig (1997) accepted this synonym.
However, these authors do not discuss some important feature s to distinguish these two species,
namely the cilia and the maculation. Based on the types here studied, they are clearly distinct
taxa, and the specimens identified by these authors should be revisited to clarify their identity.
Also the record from Vareschi (1962) to Venezuela need to be checked, since judging by the
published picture it seems to have reticular maculae and capitate soralia growing in lacinules,
typical of the species formerly placed unáer Rimelia.
The holotype collection is constituted by two larger well-preserved thalli and one smaller (5
cm broad), not as characteristic as the others. Apothecia are absent. Hale (1965) gave a
description of apothecia he found probably in some additional specimen examined (see above in
the description).
Distribution: Africa (Winnem 1975, Hale 1965, Krog & Swinscow 1981) Ásia (Kurokawa &
Lai 2001) and Américas (Hue 1899, Hale 1965, Nash & Elix 2002a). In South America, it is
known to Argentina, Brazil (Hale 1965) and Venezuela (Vareschi 1962, 1973). In Brazil, it was
recorded to the States of Minas Gerais (Hale 1965), Paraná (Osório 1977b), Rio Grande do Sul
(Canêz 2005) and São Paulo (Hale 1965).
50
Parmotrema lichexanthonicum Elias aro & Adler
Mycotaxon63:49. 1997.
Type: Brazil, Minas Gerais State, Municipality of Santana do Riacho, Serra do Cipó, Alto
Palácio, 19°22'S, 43°32'W, saxicolous, in "campo rupestre", 01.11.1989, leg. S. Eliasaro 807
(holotype: UPCB-24699!; isotypes: BAFC-37875! and CANB).
(Fig. 35)
Holotype description
Thalhis brownish (herbarium), lobate, loosely adnate, saxicolous, 13 cm broad. Lobes irregularly
branched, somewhat crowded, 5-15 mm wide, surface continuous or inconspicuously reticulate-
cracked in some lobes, usually smooth, lustrous to opaque, becoming slightly cracked in the
older lobes; apical zone more or less rounded to irregular; margin broadly crenate, sinuous,
undulated. Maculae weak, effigurate or assuming a pseudo -reticular pattern, more visible in the
amphithecia. Cilia black, simple or more usually cespitose, short 0.2-0.6 (-1.0) x 0.01-0.05 mm,
frequent, friable (sparse for almost ali upper surface). Lacinules, pustules, isidia and soredia
absent. Medalla white. Undersurface black, lustrous, strongly papillate, with cracks, rarely
veined; marginal zone absent to black or dark brown, lustrous, 1-3 mm wide, naked, with an
attenuated limit, smooth or papillate; rhizines black, simple or rarely furcated, although
sometimes difficult to ascertaining since usually the rhizines are glued in tufts by their ápices,
frequently short, 0.1-0.6 (-1.0) x 0.01-0.04 mm, very abundant, evenly distributed.
Apothecia cupuliform, 1-10 mm in diameter, stipitate, submarginal to laminai, margin smooth
or sometimes deeply dentate, amphithecium maculate, rugose, disc dark brown, epruinose,
perforate; ascospores slightly ovoid to ellipsoid, 14-16.5 x 6.5-7.5 um, episporium ca. 1 um.
Pycnidia laminai to more usually submarginal, conspicuous, with or without prominent margin,
abundant, ostiole black; conidia short- filiform, 6.5-9.0 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV+ yellow.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), lichexanthone (minor).
Remarks: Parmotrema lichexanthonicum is characterized by the presence cespitose cilia,
medulla with lichexanthone and salazinic acid, and the absence of vegetative propagules.
The presence of cespitose cilia in this species is notorious and was already pointed out by
Ribeiro (1998). It is correlated with the presence of lichexanthone in the medulla. Similar species
with salazinic acid and lichexanthone include P. pontagrossense and P. spinibarbe (sorediate),
and P. idtralucens (isidiate).
While identifying specimens of Parmotrema lichexanthonicum, we realized that the medulla
not always shows an evident yellow bright under the UV lamp. Donha (2005) noticed the same
for P. pontagrossense and suggested the TLC analysis as essential to assure the presence of
lichexanthone. However, the experience has demonstrated that the presence of cespitose cilia
together the typical dark reddish brown shiny marginal zone are strong indication of the medullar
lichexanthone presence in Parmotrema species.
Both the holotype and isotype are well developed thalli, and the isotype (BAFC) agrees well
with the holotype in ali features. Indeed, the faint maculate upper surface is more easily seen in
this thallus.
An important and relatively recent problem is the exploration of Brazilian lichens in a
commercial scale. Christensen & Sipman (1998) noted that Brazilian lichens were sold in
European markets. They thought that one of the species was Parmotrema cetratam (A eh.) Hale.
51
However, cio se examination of the specimens deposited in C proved they are actually
Parmotrema lichexanthonicam . So, it is possible that this apparently endemic lichen be an
endangered species.
Distribution: known to Brazil, where it was recorded to the States of Minas Gerais (Eliasaro &
Adler 1997, Ribeiro 1998) and São Paulo (Ribeiro 1998).
Parmotrema livido-tesselatum (Hue) Spielmann & Marcelli, comb. nov.
Parmelia livido-tesselata Hue, Nouvelles Archives du Muséum d'Histoire Naturelle de Paris, sér.
4 (1): 191, plate V, fig. 2. - Type: Uruguay, Montevideo, Cerro Melones, saxicolous, June 18,
1876, leg. Fruchart s.n. (lectotype: P!; isotypes: TUR!).
(Fig. 36)
Holotype description
Thallus brownish (herbarium), lo bate, loosely adnate, saxicolous and over mosses, up to 27 cm
broad (combining the parts). Lobes irregularly branched, somewhat crowded, growing in
multiple directions (i.e., not radial), 5-15 mm wide, surface continuous to reticulate-cracked,
opaque to lustrous, strongly maculate, pruinose in some lobe ápices; apical zone rounded; margin
sublacinulate, with well marked incisions. Maculae strong, reticular, sometimes extensive
(hypermaculate), originating cracks. Cilia black, simple to furcated, rarely more branched,
sometimes thickened, 1.0-3.5 x 0.05-0.20 mm, abundant. Soredia farino se, initially coming
from the margins of the areolae, coalescing with the age and forming plane or extensive, laminai
or more commonly submarginal soralia, sometimes with a pustular appearance. Lacinules
simple, projected from the margin of the lobes, plane, apex rounded/to ± truncate, up to 5 mm
long, 0.5-2.0 mm wide. Isidia absent. Medulla white. Under surface black, lustrous, usually
papillate, less commonly smooth or rugose, with cracks; marginal zone dark brown, lustrous,
1.5-4.0 mm wide, naked, with an attenuated limit, usually smooth, rarely slightly papillate;
rhizines black, simple or sometimes squarrose or furcated, 0.2-2.5 x 0.02-0.10 mm, abundant,
irregularly distributed.
Apothecia concave, 5-6 mm in diameter, substipitate, laminai, margin incised (difficult to
define since the apothecia are not in good conditions), amphithecium scrobiculate, disc brown,
epruinose, apparently perforate; ascospores ellipsoid 13 x 6.5-7.5 um, episporium ca. 1 um (see
the notes below). Pycnidia not found (several black structures investigated, but without conidia).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC (J.A. Elix, 25.11.1996, label with the isotype): atranorin (major),
chloroatranorin (minor), salazinic acid (major), consalazinic acid (minor).
Remarks: Parmotrema livido-tesselatum has a wide and distinct bare zone on the underside,
thickened, tortuous, tapered and usually furcated cilia, an upper surface strongly maculate, and
the soralia often laminai and eruptive rose from the thallus cracks.
Hale & Fletcher (1990) situated this species as synonym ofRimelia reticulata. Actually, this
lichen is more close to Parmotrema austromaculatum, differing by the presence of soredia. It is
also interesting to note that, in P. livido-tesselatum, the sorediate lobes do not become involute,
as commonly happen to other "rimelioid" species.
52
The holotype collection (PC) is composed of four thalli, or possibly parts of just one thallus,
since Hue's (1899) thallus measure is 27 cm broad, which agree with our observation. One of the
parts contains two smashed apothecia, and is glued to a paper with the annotation "n. 33".
Apparently Hue saw just one apothecia, the more developed, and we chose the second one to
take the ascospores measures, as to preserve the main apothecium. The hymenium is not well
developed, and just two ascospores were found. They are 13 x 6.5-7.5, so matching the variation
described in the protologue (12-16 x 7-8 um).
There are two lichens preserved in TUR, numbered "02620" and "02621", which were part of
the Herbarium Vainio. They are very similar in every feature with the lectotype from PC, and the
labels are indicating "isotype" In fact, it seems clearly that they are parts of the collection from
PC. However, Hue's (1899) original publication cited two collections: one from Uruguay and
another from "Africa meridionalis". Probably this information led Hale & Fletcher (1990) to
choose the specimen from PC as lectotype, and the specimen from TUR (probably only the n.
02620, since it is the only one with Hale annotations) as "isolectotype". This term is not used by
the current ICBN (Vienna Code), and so it can be called "duplicate of the lectotype".
The specimen referred by Hue (1899) to South Africa requires to be revised, since it can
indicate an interesting distribution pattern.
Distribution: Uruguay and South Africa (Hue 1899, Zahlbruckner 1930a).
Parmotrema magnum (Lynge) Spielmann & Marcelli, comb. nov.
Parmelia magna Lynge, Arkiv for Botanik 13 (13): 83. 1914. - Type: Brazil, Minas Gerais
State, Municipality of São João d'el Rey [dei Rei], 31. VIII. 1892, leg. G.O.A. Malme 269
(holotype: S!; isotypes: LD!, MO, UPS!).
= Parmelia microdactyla Hale, Contributions from the United States National Herbarium 36 (1):
21. 1960. - Type: Brazil, [Rio de Janeiro State], Municipality of Rio de Janeiro, "Pâ õversta
toppen av Corcovado" [at the very top of the Corcovado mountain], leg. Fr. Widgren
(holotype: UPS!; isotype: S!).
(Figs. 37-40)
Holotype description
Thallus yellowish, lobate, loosely adnate, probably saxicolous, 17 cm broad. Lobes irregularly
branched, laterally overlapped to crowded, 5-15 mm wide, surface continuous, smooth,
sublustrous, becoming slightly rugose towards the center; apical zone rounded; margin crenate to
sinuous, undulated. Maculae absent. Cilia black, simple or furcated, up to 1.5 mm long, 0.05-
0.10 mm wide, few. Lacinules, pustules, soredia and isidia absent. Medulla white. Undersurface
black, sublustrous, smooth to rugulose, papillate near the margins, or rarely with veins, with
cracks; marginal zone dark brown, lustrous, 1.0-3.5 mm wide, naked, with an attenuated limit,
smooth or rugulose; rhizines black, simple or rarely irregularly branched, delicate, up to 2.5 mm
long, 0.02-0.06 mm wide, abundant, distributed in groups.
Apothecia ílattened or cupuliform, usually lacerate and with the margins involute, up to
10 mm in diameter, short stipitate, submarginal, margin smooth or incise, amphithecium
maculate, rugose or scrobiculate at maturity, disc brown, epruinose, imperforate; ascospores
ellipsoid, 10.0-12.5 x 7.5-8.0 um, episporium ca. 1 um. Pycnidia submarginal, conspicuous,
with or without prominent margin, frequent, ostiole black; conidia short-filiform, 6.0-7.5 x ca.
1.0 um.
53
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C+ rose, KC+ red, P+
strong yellow, UV-.
TLC in solvent C, HPLC (J.A. Elix, 10.V.1996, label with the holotype in S): usnic acid (minor),
salazinic acid (major), consalazinic acid (minor), gyrophoric acid (major), lecanoric acid (trace).
Remarks: Parmotrema magnum can be recognized by the yellowish large thallus (up to 17 cm)
and wide lobes (5-15 mm), with not abundant cilia up to 1.5 mm long. The apothecia are usually
lacerate, with a rugose to scrobiculate amphithecia (Fig. 38). The medulla reacts C+ rose, KC+
red, an indicative of the abundance of gyrophoric acid.
It is close to P. delicatulum, from which it was already considered a synonym (Hale 1965).
However, P. delicatulum has a smaller sublaciniate thallus with narrow lobes (1.0-3.5 mm), cilia
abundant, simple and longer (up to 2.8 mm), and smooth entire amphithecia. Although the
gyrophoric acid was reported to this species (see above), it is not abundant enough to cause a
positive spot test, and so the medulla is C-, KC-.
These differences are being considered here as important to maintain these taxa as good
species. Future studies with additional specimens from a wider área can corroborate or not this
positioning.
A special problem occurs with Parmelia microdactyla Hale. This species was proposed (Hale
1960) based on the presence of marginal lobules and very sparse cilia. Later on, Hale (1965)
assumed this difference as a morphological variation, and situatedP. microdactyla as a synonym
of P. delicatula (together with P. magna). The holotype is represented by a well-preserved
thallus 8.5 cm broad (Fig. 39), with pruinose lobes (at the ápices), abundant lobules (Fig. 40) and
rare, short cilia (up to 0.5 mm long). It is also very close to Parmotrema flavescens, whose
isotype (S) was found growing together. However, no sign of isidia was found. The chemistry of
the isotype was made through TLC in solvent C and HPLC (J.A. Elix, 10.V.1996), and revealed
usnic acid (minor), salazinic acid (major), consalazinic acid (minor), gyrophoric acid (major),
lecanoric acid (trace). The medulla of this specimen is C-, KC+ rose, and so intermediary
betweenP. delicatula andP. magna.
Unfortunately, apothecia are unknown both in Parmotrema microdactyla and P. flavescens,
and so additional differences or similarities could not be compared. By the moment, we place P.
microdactyla underP. magnum because the almost absence of cilia.
Distribution: known for Brazil, being recorded to the States of Minas Gerais (Lynge 1914, Hale
1 960) and Rio de Janeiro (Hale 1 960).
Parmotrema mantiqueirense Hale
Bibliotheca Lichenologica 38: 113. 1990.
Type: Brazil, Minas Gerais State, Serra da Mantiqueira, Oberhalb von Vila Monte Verde,
zwischen 1880 m und der Gipfelregion des Pedra Redonda in 2000 m, 3.VII.1979, leg. K. Kalb
s.n. (holotype: Kalb herbarium; isotype: US!).
(Fig. 41)
Isotype description
Thallus whitish gray, lobate, loosely adnate, over rocks with mosses and plant debris, 11.5 cm
broad. Lobes irregularly branched, laterally overlapped to quite crowded, 7-20 mm wide, surface
continuous, smooth, lustrous, becoming irregularly scared and cracked towards the center; apical
zone ± rounded; margin broadly crenate to sublacinulate, undulated. Maculae absent. Cilia
54
black, simple to furcated, up to 2.5 mm long, often thickened, 0.10-0.15 mm wide, frequent.
Lacinules, pustules, soredia and isidia absent. Medulla white. Undersurface black, lustrous,
smooth, rugulose, papillate or rarely veined, with cracks; marginal zone brown, lustrous, 4-
9 mm wide, naked or in part rhizinate, with an attenuated limit, smooth to slightly rugose or with
rhizinal papillae; rhizines black, simple or rarely furcated, up to 4 mm long, 0.02-0.10 mm wide,
abundant, evenly distributed.
Apothecia absent [according to Eliasaro (2001), apothecia laminai, adnate, 2-5 mm in
diameter, margin smooth, disc brown, imperforate; ase osp ores 13-18 x 6-10 um]. Pycnidia
laminai or more often submarginal, conspicuous, with or without prominent margin, abundant,
ostiole black; conidia filiform, 8-12 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema mantiqueirense is distinguished by the large, conspicuously ciliate lobes,
filiform conidia and the absence of propagules.
It is close to P. latissimum, an eciliate species with sublageniform conidia.
Apothecia are absent both in the holotype (Hale 1990) and lectotype. However, Eliasaro
(2001 ) found one fertile specimen from Paraná State, Brazil (see above in the description).
In P. mantiqueirense some rhizines can oceupy part of the marginal zone, mainly in the form
of "rhizinal papillae" (Awasthi 1976) or "rhizines in development" (Fleig 1997). This feature
was already noted to Parmotrema internexum (Nyl.) Hale ex DePriest & Hale (Spielmann &
Marcelli 2009).
Distribution: South America, where it was reported to Brazil (Marcelli 2008). In Brazil it was
recorded to the States o f Minas Gerais (Hale 1990, Fleig 1997, Ribeiro 1998), Paraná (Fleig
1997; Eliasaro 2001) and Rio Grande do Sul (Canêz 2005, Spielmann 2005, 2006).
Parmotrema margaritatum (Hue) Hale
Phytologia28 (4): 337. 1974.
Parmelia margaritata Hue, Nouvelles Archives du Muséum d'Histoire Naturelle de Paris ser. 4
(1): 193. 1899. - Type: U.S.A, Ohio, corticicolous, leg. Sullivant (lectotype: P!).
(Fig. 42)
Holotype description
Thallus brownish (herbarium), sublaciniate, loosely adnate, substrate, 6-17 cm broad.
Sublaciniae crowded, 5-15 mm wide, surface continuous, smooth, opaque to sublustrous,
becoming cracked towards the center; apical zone lacinulate, undulated and irregular. Maculae
effigurate or sometimes punctiform, often joined and covering mo st of the thallus surface
(hypermaculate), and so quite difficult to realize at first sight, laminai, not originating cracks.
Cilia black, simple, up to 2 mm long, usually thickened, 0.04-0.12 mm wide, frequent. Lacinules
abundant, simple, dichotomous or irregularly branched, plane, apex usually acute, sometimes
truncate, 2.0-10.0 x 0.5-3.0 mm. Soralia whitish, orbicular, subapical on the lacinules, which
can become involute as the soralia grows and become coníluent; soredia farino se to subgranular.
Pustules and isidia absent. Medulla white. Undersurface black, lustrous, smooth to slightly
55
rugose or papillate, rarely veined, with abundant cracks; marginal zone dark brown, lustrous, 1-
5 mm wide, naked, with an attenuated limit, smooth, slightly rugose or papillate; rhizines black,
monomorphic, up to 2.5 mm long, 0.01-0.40 mm wide, simple or irregularly branched,
especially near the ápices, few, distributed in groups.
Apothecia unknown. Pycnidia submarginal to laminai and restricted to the lacinules,
conspicuous, usually with prominent margin, abundant, ostiole black; conidia filiform, 10-15 (-
18) x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema margaritatum is distinguished by the abundantly lacinulate thallus, the
soralia orbicular or coníluent growing on the lacinules, the effigurate maculae usually becoming
hypermaculate, the thickened and simple cilia and the black undersurface, with a dark brown and
erhizinate marginal zone.
Also sorediate, P. petropoliense has pustuloid raised structures that give rise to soredia, long
(up to 3 mm) and frequently furcated cilia, brown undersurface with dichotomous rhizines and
effigurate maculae.
The holotype collection is composed by six thalli. Three of them are glued in a paper, in one
sheet (PC0 116176) and three are not glued, in other sheet (PC0 116177). Both have full
indications of collector, Hue's handwriting, etc, and actually, they are very similar in ali
features. Hale (1965) illustrated only part of the larger specimen from the sheet PC01 16177, and
so if a lectotype must be chosen, this is probably the best choice, since the undersurface can also
be adequately studied.
Distribution: Ásia (Awasthi 1976, Chen et ai 2005, Divakar & Upreti 2005), North America
(Hale 1965, Taylor 1967, Brodo et ai. 2001) and South America, where it was reported to
Venezuela (Feuerer 2008) and the State of Rio Grande do Sul in Brazil (Canêz 2005).
Parmotrema masonii Ferraro
Hickenia 1 (34): 191. 1979.
Type: Argentina, Prov. Comentes, Depto. Concepción, Paso Crucecita, 20-VI-1974, leg. L.I.
Ferraro 215 (holotype: CTES!; isotype: US!; paratype: CTES!).
(Fig. 43)
Holotype description
Thallus yellowish, lobate, loosely adnate, corticicolous, 10 cm broad. Lobes irregularly
branched, laterally overlapped, 2-10 mm wide, surface continuous, smooth, opaque, becoming
strongly rugose and cracked towards the center; apical zone rounded; margin crenate, sometimes
with deep sinuses, undulated. Maculae indistinct, punctiform, laminai. Cilia black, simple,
usually tapered and turning upwards, up to 0.7 mm long, 0.05-0.10 mm wide, frequent.
Lacinules, pustules, soredia and isidia absent. Medulla white. Undersurface black, sublustrous,
smooth, rugose, papillate, with cracks; marginal zone dark brown, sublustrous, 1-2.5 mm wide,
naked, with an attenuated limit, smooth, rugose or papillate; rhizines black, simple or rarely
branched, up to 1.5 mm long, 0.02-0.05 (-0.10) mm wide, abundant, ± evenly distributed.
56
Apothecia cupuliform, 0.5-5.5 mm in diameter, short-stipitate, usually submarginal, margin
crenate, frequently involute, amphithecium smooth to irregular or rugose, pruinose, disc brown,
slightly pruinose, imperforate; ascospores ellipsoid to ovoid, 15-16.5 x 7.5-8.0 um, episporium
ca. 1 um. Pycnidia submarginal, conspicuous, with or without prominent margin, abundant,
ostiole black; conidia filiform, 7-12 x ca. 1.0 um.
Color tests: córtex K+ yellowish, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: usnic acid (minor), salazinic acid (major), consalazinic acid (minor), hypoconstictic
acid (minor).
Remarks: Parmotrema masonii is characterized by the short cilia (up to 0.7 mm), the presence
of salazinic and hypoconstictic acid and the absence of propagules.
Related species include P. delicatulum (longer cilia and with different chemistry), P.
flavescens (isidiate), P. mirandum, P. nylanderi andP. pectinatum (sorediate).
The holotype of P. masonii has the upper surface strongly rugose (resembling that of P.
madylinae), short and tapered cilia. The isotype agrees very well with the holotype, except that it
has slightly shorter ascospores 13.5-15 um long and apothecia frequently irregularly "coronate"
with black dots, probably formed by some parasitic fungus. The paratype received from CTES
presents several differences in the relation to the holotype and isotype: 1) larger, lacerate
apothecia (up to 8 mm in diameter); 2) smaller ascospores, 10-12.5 x 6-7.5 um, episporium 0.5-
1.0 um; 3) upper surface only slightly rugose and 4) cilia usually thinner and longer (up to 1.5
mm). Nevertheless, the conidia and overall morphology are similar.
The specimen from São Paulo (Marcelli & Barbosa 35720) shares the same chemistry of P.
masonii, but also has thinner and longer cilia (up to 1.5 mm) and the upper surface is only
slightly rugose. It does not produce apothecia but the conidia data (filiform, 8-1 1 x ca. 1 um
according to a label from P. Jungbluth dated 2005) agrees roughly with the measurements from
the holotype. Since we did not have access to a rich collection of this species, it is premature to
assign these differences to different taxa by the time.
Distribution: South America, where it was recorded to Argentina (Ferraro 1979), Brazil (Fleig
1997) and Paraguay (Fleig 1997). In Brazil it is known for the States of Rio Grande do Sul (Fleig
1 997) and São Paulo (Jungbluth 2006).
Additional specimen s examine d: Argentina, Prov. Comentes, Depto. Concepción, Paso
Crucecita, on Sapium sp. (Euphorbiaceae), 20-VI-1974, leg. L.I. Ferraro 206 (CTES-78596).
Brazil, São Paulo State, Municipality of São Manuel, Fazenda Palmeira da Serra, private cerrado
protected área, on tree trunk in the woods edge, 03.VI.2003, leg. M.P. Marcelli & S.B. Barbosa
35720 (SP).
Parmotrema maximum (Hue) Spielmann & Marcelli, comb. nov.
Parmelia máxima Hue, Nouvelles Archives du Muséum d'Histoire Naturelle de Paris, ser. 4 (1):
193. 1899. - Type: México, Chimaleapa n, Cerro de Santiago, near Lerma, apparently
saxicolous, 26.VI.1890, leg. Maury 3318 (holotype: P!).
(Fig. 44)
57
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, probably saxicolous, 13.5-17 cm broad.
Lobes irregularly branched, laterally overlapped to ± crowded, 0.5-25 mm wide, surface
continuous, smooth or with rounded tiny depressions, lustrous, becoming irregularly cracked
towards the center; apical zone rounded; margin broadly crenate to lacinulate, undulated to
sinuous. Maculae absent or irregular and conspicuous in some lobes. Cilia black, simple to
furcated or dichotomous, up to 2.5 mm long, usually thickened, 0.04-0.10 mm wide, frequent.
Lacinules simple to dichotomous, plane, apex usually rounded, sometimes truncate or acute, up
to 5 mm long, 0.5-5.0 mm wide. Pustules and isidia absent. Soralia linear continuous or
sometimes growing backwards or becoming labriform, marginal and on the lacinules ápices;
soredia farinose to granular. Mediãla white. Undersurface black, lustrous, smooth to rugose or
papillate; marginal zone absent (black) or brown to dark brown, lustrous, 2-6 mm wide, naked,
with an attenuated limit, smooth, slightly rugose or papillate, with cracks; rhizines black, usually
simple, less frequently irregularly branched, up to 3 (-4) mm long, 0.01-0.10 (-0.20) mm wide,
abundant, irregularly distributed.
Apothecia plane (pressed down), 1-2.3 cm in diameter, apparently substipitate, margin
smooth to incised or lacerate, amphithecium maculate, with irregular depressions or rugose, disc
dark brown, epruinose, perforate or not; ascospores ellipsoid, 12.5-15 x 7-9 um, episporium ca.
1 um. Pycnidia submarginal, little conspicuous, without prominent margin, few, ostiole black;
conidia short-filiform, 7-10 x ca. 1 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema maximum is charactenzed by the presence of marginal soralia, occurring
also at the ápices of the lacinules, the broad nude under margin, the size of ascospores and the
short-filiform conidia.
This species was up to know include in the synonymy of P. stuppeum, which however do not
develop lacinules and additionally has sublageniform conidia (Nash & Elix 2002b). More similar
morphologically is P. margaritatum (see Hue 1899, Hale 1979), however distinct by the
effigurate maculae.
The holotype is composed by two thalli, one of them bearing several apothecia and labeled
"32". It was clearly cut off at the middle. The second thallus is quite similar to the first, however
no apothecia are present, and it is not part of the first thallus. Morphologically and chemically
the two thalli agree well.
Distribution: México (Hue 1899).
58
Parmotrema mirandum (Hale) Hale
Phytologia28 (4): 337. 1974.
Parmelia miranda Hale, Contributions from the United States National Herbarium 36 (5): 273.
1965. - Type: México, State of Oaxaca, northwest of Tehuantepec, Km 686 on highway 190,
Pine-scrub oak mountainside, on deciduous trees, elev. 1160 m, 31. III. 1960, leg. M.E. Hale &
T.R. Soderstrom 20653 (holotype: US!; isotypes: COLO!, DUKE!, LISU, MSC!, REN, S!,
TNS!).
(Fig. 45)
Holotype description
Thalhis yellowish, lobate, ± adnate, corticicolous, 5.5-1 1.0 cm broad. Lobes irregularly
branched, laterally overlapped to ± crowded in the center, 4-9 mm wide, surface continuous,
smooth to slightly scrobiculate or rugose, opaque, becoming irregularly cracked in some áreas;
apical zone rounded, often pruinose; margin broadly crenate, sometimes more incised, undulated.
Maculae inconspicuous, irregular. Cilia black, simple or rarely furcated, 0.2-1.0 (-1.5) x 0.03-
0.05 mm, frequent. Sor alia marginal and usually extensive, also in the ápices of the sublacinules,
turning the lobe ápices involute; soredia subgranular. Lacinules, pustules and isidia absent.
Medulla white or with red stains in some áreas (salazinic acid in decomposition). Undersurface
black, lustrous, rugose, papillate or rarely veined, with cracks; marginal zone brown or
sometimes pale brown or with beige stains, lustrous, \-4 mm wide, naked, with a sharp limit,
smooth, rugose or papillate; rhizines black, simple, 0.2-1.5 x 0.01-0.05 mm, abundant, evenly
distributed in the distai parts but absent in large áreas at the center.
Apothecia unknown. Pycnidia (from the MSC isotype) submarginal, conspicuous, with
prominent margin, rare, ostiole black; conidia short-filiform, 6-9 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC: usnic acid (minor), salazinic acid (major), consalazinic acid (minor),
hypoconstictic acid (minor).
Remarks: Parmotrema mirandum is recognized by the yellowish (usnic acid) sorediate thallus
with salazinic acid and hypoconstictic acid. The disco very of hypoconstictic acid in this species
places it close to Parmotrema masonii, suggesting it as the sorediate pair.
The holotype consists of three well developed thalli, and by the great abundance of isotypes
(some also with two or three thalli), one can assume the species was quite abundant in the type
locality.
About 20 supposed pycnidia (it is difficult to prove if they will produce conidia or not, since
they are actually only black dots in the surface, and at the microscope no special structure can be
seen) were examined from the holotype; however, no conidia were found. Only one "true"
pycnidium was found in the isotype from MSC, from where the measurements were obtained.
The examined isotypes are similar in the overall morphology to the holotype, but some
particularities can be pointed: the isotype from TNS has more distinct beige marginal zones in
the undersurface, and they seem to be clearly associated with the lobes that produce soralia. The
same is true to the isotype from MSC, which additionally has wider lobes, up to 14 mm wide.
Large lobes (equally up to 14 mm) were found in the isotype from S. Finally, the isotype from
DUKE develops some strongly rugose pustulate-like áreas in the thallus center.
59
Distribution: North America (Hale 1965), and South America, where it was recorded to Brazil
and Venezuela (Feuerer 2008). In Brazil it is known for the States of Minas Gerais (Ribeiro
1998), Paraná (Eliasaro 2001) and São Paulo (Ribeiro 1998).
Parmotrema neotropicum Kurok.
Mycotaxon 5 (2): 437. 1977.
Type: México, State of Chiapas, Pine-oak forest on mountainside, 2120 m alt., Km. 1145 on
highway 190, west of San Cristóbal, on conifers, 25.111.1960, leg. M.E. Hale & T.R. Soderstrom
20190 (holotype:US!).
= Rimeliella neotropica (Kurok.) Kurok., Armais of the Tsukuba Botanical Garden 10: 6. 1991 .
= Canomaculina neotropica (Kurok.) Elix, Mycotaxon 65: 477. 1997.
(Fig. 46)
Holotype description
Thallus buff yellowish, lobate, adnate to loosely adnate, corticicolous, 9.5 cm broad. Lobes
irregularly branched, ± crowded, 3-12 mm wide, surface continuous, with rounded depressions
or cracked, smooth, sublustrous, becoming rugose and cracked in the center; apical zone
rounded; margin crenate-incised, undulated. Maculae weak, effigurate, laminai, not originating
cracks. Cilia black, simple or rarely furcated, up to 1 .2 mm long, 0.02-0. 10 mm wide, few. Isidia
concolor to the thallus or paler, apex usually dark brown, simple to coralloid, up to 1.0 mm high,
0.05-0.50 mm wide, erect, firm or breaking into soredioid granules, sometimes with lateral cilia,
laminai or more usually submarginal to marginal. Lacinules, pustides and soredia absent.
Medulla white. Undersurface dark brown to brown, lustrous or opaque, abundantly veined,
without cracks; marginal zone absent; rhizines black, dimorphic, the shorter ones simple and
usually curled, up to 1 mm long, 0.01-0.05 mm wide, abundant, evenly distributed; the longer
ones up to 4 mm long, 0.05-0.15 mm, usually irregularly branched, especially near the apex,
few, distributed in groups.
Apothecia unknown. Pycnidia submarginal, conspicuous, without prominent margin, rare,
ostiole black; conidia filiform, 7-1 1 x ca. 1 .0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), usnic acid (minor), isousnic acid (minor),
salazinic acid (major), consalazinic acid (minor).
Remarks: Parmotrema neotropicum is distinguished by the isidiate thallus, the isidia sometimes
breaking up in soredioid granules, the effigurate maculae, the abundantly veined, brown
undersurface with dimorphic rhizines and the presence of usnic acid.
It can be confused with P. subtinctorium, but in this species the isidia remain entire, the
undersurface is only slightly rugose to papillate, the usnic acid is absent, and produces
norlobaridone in the medulla.
The holotype is composed by two distinct thalli (i.e., not parts of the same thallus) that agree
with each other in ali important features.
Distribution: Brazil (Marcelli 2008), Cuba, México, U.S.A (Hale 1977, Kurokawa 1991). In
Brazil it was reported to the States of Goiás (Kurokawa 1991), Minas Gerais (Ribeiro 1998),
60
Paraná (Donha 2005), Pernambuco, Rio de Janeiro (Kurokawa 1991), Rio Grande do Sul (Fleig
1997) and São Paulo (Marcelli 1992, Marcelli & Benatti 2008).
Parmotrema nudum (Hue) Spielmann & Marcelli, comb. et stat. nov.
Parmelia reticidata f nuda Hue, Nouvelles Archives du Muséum d'Histoire Naturelle de Paris,
sér. 4 (1): 177, plate V, fig. 2. - Type: Uruguay, prope Montevideo, Cerro Melones, saxicolous,
1875 and 1876, leg. Fruchart s.n. (lectotype: P!).
(Fig. 47)
Holotype description
Thallus brownish, sublaciniate, apparently loosely adnate, saxicolous, 6-11 cm broad;
sublaciniae irregularly branched, crowded, (1-) 2-6 mm wide, surface continuous, reticulate,
irregularly cracked towards the centre, smooth to strongly rugose and forming pustules; apical
zone truncate or slightly rounded, sometimes acute; margin usually broadly sinuous, plane to
undulated. Maculae distinct, reticular, laminai, originating cracks that take part in the formation
of the pustules; Cilia black, simple, 0.2-0.9 mm, abundant. Pustules rugose to shapeless,
laminai, remaining entire, often formed in the areolae contact. Lacinules, soredia and isidia
absent. Isidia absent. Medulla white. Undersurface not well studied [thallus pasted in a paper
card, description based on just a small part of one thallus] lustrous, smooth to rugose, with
cracks; marginal zone dark brown, lustrous, up to 4 mm wide, rhizinate, with an attenuated limit,
smooth or papillate; rhizines black, usually simple, up to 1 mm long, 0.01-0.05 mm wide,
abundant, more or less evenly distributed.
Apothecia sometimes plane (probably pressed down) to more commonly concave, cupuliform
or urceolate, up to 7 mm in diameter, substipitate, laminai, margin smooth to strongly pustulate,
sometimes lacerate, amphithecium strongly maculate near the border, smooth, rugose (at times
strongly) or pustulate, disc dark to light brown, epruinose, imperforate or rarely perforate at
maturity; ascospores ellipsoid to oblong, 12-14 x 7.5-9.0 um, episporium ca. 1 um. Pycnidia
mainly submarginal, conspicuous, usually with prominent margin, abundant, ostiole black;
conidia filiform 9.0-12.5 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema nudum can be distinguished by the reticular maculae and laminai
pustules.
This species was up to know considered a synonym of P. reticulatum (Hale & Fletcher 1990).
However, the pustular upper surface and absence of soredia are sufficient features to distinguish
the se two species easily.
It can be more properly compared with P. granulare, which has abundant lacinules but
produces soredia, and P. ruminatum, an eciliate species where the pustules sometimes ílakes off
the areolae.
The lectotype is well-preserved and developed, although glued to a paper, so that the
underside could not be completely described.
Distribution: Uruguay (Hue 1899).
61
Parmotrema nylanderi (Lynge) Hale
Smithsonian Contributions to Botany 33: 54. 1976.
Parmelia nylanderi Lynge, Arkiv for Botanik 13 (13): 82. 1914. - Type: Brazil, Mato Grosso,
Serra da Chapada [nowadays Municipality of Chapada dos Guimarães], near São Jeronymo,
supra rupes apricas, 03.VI.1894, leg. G.O.A. Malme 2747 (holotype: S!).
= Parmelina nylanderi (Lynge) Hale, Phytologia 28 (5): 483. 1974.
(Fig. 48)
Holotype description
Thallus yellowish, lobate, loosely adnate, saxicolous, 6.0-7.5 cm broad. Lobes irregularly
branched, laterally overlapped, 1.0-4.5 mm wide, surface continuous or cracked, smooth,
lustrous in the young regions, becoming dull rugose and cracked towards the center; apical zone
rounded; margin crenate, marked by a distinct black line, sometimes sublacinulate, undulated.
Maculae absent. Cilia black, simple or rarely bifurcated near the apex, usually short, and
sometimes thickened, 0.2-1.0 (-1.5) x 0.05-0.10 mm, frequent, although in some lobes totally
absent. Pustides rugose or orbicular, or multiform, submarginal extensive, usually arranged in
groups and advancing over the lamina, originating granular soredia. Lacinules and isidia absent.
Mediãla white. Undersurface black, lustrous, smooth or rarely rugose, with cracks; marginal
zone brown, lustrous, 1-2 mm wide, naked, with a sharp or attenuated limit, usually smooth, less
frequently slightly rugose or papillate; rhizines black, simple, furcated or less frequently
dichotomous, up to 2 mm long, thin to thickened, 0.02-0.10 mm wide, abundant, irregularly
distributed.
Apothecia unknown. Pycnidia submarginal, inconspicuous, without prominent margin, few,
ostiole black, immature; conidia not found (ca. 20 pycnidia examined).
Color tests: córtex K-, UV-; medulla K+ yellow — > blood red, C+ rose, KC+ reddish rose, P+
strong yellow, UV-.
Chemistry: usnic acid (minor), salazinic acid (major), consalazinic acid (minor), hypoconstictic
acid (minor), gyrophoric acid (minor).
Remarks: Parmotrema nylanderi is distinguished by the lobate yellowish thallus (usnic acid),
the presence of salazinic acid and gyrophoric acid in the medulla (K+ yellow — > red, C+ rose)
and the production of pustules.
Another close species, which also produces pustular soredia, is Parmotrema pectinatum. It
differs by the laciniate thallus with narro w laciniae (0.8-3.0 mm) copiously ciliate at the
margins. Apothecia and conidia are unknown, however, and since they are also unknown in P.
nylanderi, further collections are necessary to establish more concretely the relations of these
two species.
Another species with salazinic acid and usnic acid do not have gyrophoric acid: P. mirandum
(sorediate), P. delicatulum andP. masonii (without propagules) and P. flavescens (isidiate).
The holotype of P nylanderi is constituted by four pieces, and could well be part of the same
thallus. In the protologue (Lynge 1914) the thallus size is 6-7 cm. But this lichen was collected
by Malme several years earlier, inl894, and could well have cracked before the description.
Although Hale (1960) first defined the specimen Malme 21 Al as holotype, later on (Hale
1976) he considered it to be a lectotype. However, since it is the only specimen recorded in the
protologue (Lynge 1914), and as apparently there are no additional types, it must be categorized
as holotype, according to the Article 9.1 Note 1 of the Vienna Code (McNeill et ai. 2007).
62
Distribution: known only for Brazil, where it was recorded to the States of Minas Gerais
(Ribeiro 1998), Mato Grosso (Lynge 1914, Hale 1960) and São Paulo (Hale 1976, Jungbluth
2006). The record from Ribeiro (1998) needs to be reinvestigated, since it has negative thallus
reactions (Jungbluth 2006).
Parmotrema parahypotropum (W.L. Culb.) Hale
Phytologia 28 (4): 338. 1974.
Parmelia parahypotropa W.L. Culb., The Bryologist 76 (1): 29. 1973. - Type: Taiwan
(Formosa), Kural, 250 malt, 01.11.1964, leg. Kurokawa 1487 (holotype: TNS!, isotype: DUKE!).
(Figs. 49-50)
Holotype description
Thallus pale brownish (herbarium), sublaciniate?, extensively lacinulate, loosely adnate, 5.5 cm
broad. Sublaciniae dichotomous or irregularly branched, laterally overlapped to ± crowded, 1.5-
5.0 mm wide, surface continuous or sometimes with irregular depressions, smooth, opaque,
becoming slightly rugose and cracked towards the center; apical zone rounded when without
lacinules; margin sinuous-incised or more commonly lacinulate, undulated. Maculae distinct,
effigurate, laminai, rarely originating cracks. Cilia black, simple or furcated, up to 1.5 mm long,
0.03-0.08 mm wide, frequent. Lacinules abundant, usually dichotomous, plane, apex rounded,
truncate or acute, 1.0-6.0 x 0.5-2.0 mm. Soralia white, growing from the margin of the lobes
and becoming extensive towards the undersurface, the sorediate lobes becoming crescent-shaped
and crisped; soredia farinose to subgranular. Pustules and isidia absent. Medulla white.
Undersurface black to dark brown in some lobes, beige under the lacinules (probably white in
fresh material), lustrous, smooth or with reticulate veins, with few cracks; marginal zone brown
or beige under the lacinules, lustrous, 1-3 mm wide, naked, with an attenuated limit, usually
slightly rugose or veined; rhizines black or rarely with pale brown ápices, simple, usually glued
to each other, up to 1 mm long, 0.02-0.10 mm wide, abundant, irregularly distributed.
Apothecia unknown. Pycnidia submarginal to laminai, conspicuous, with or without
prominent margin, few, ostiole black; conidia unciform, 4-5 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > slowly red-orange, C-, KC-, P+
strong yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (minor), salazinic acid (minor), norstictic acid
(major), galbinic acid (major), consalazinic acid (trace).
Remarks: Parmotrema parahypotropum is characterized by the lacinulate thallus with marginal
soralia, that grow towards the undersurface, the effigurate maculae, unciform conidia and
complex medullar chemistry, with salazinic acid, norstictic acid and galbinic acid .
Soralia ontogeny is very similar to that found in P. subsumptum, the sorediate lacinules
usually becoming crescent-shaped or crisped. But the abundant lacinules, chemistry, conidia
type, furcated cilia and absence of dimorphous rhizines can be used to set these species apart.
P. parahypotropum is the only known species with salazinic acid and unciform conidia, this
last feature here being reported for the first time.
Distribution: Africa (Krog & Swinscow 1981, Kurokawa 2003, Aptroot 2008), Ásia (Culberson
1973) and Austrália (Elix 1994).
63
Parmotrema paramoreliense^N .\u. Culb. & CF. Culb.
TheBryologist84(3): 311. 1981.
Type: México, Michoacán, 33 Km E of Uruapan on the road to Patzcuaro, pine-oak woods, on a
fallen branch oíPinus sp., 28.XII.1978, leg. W.L. Ciãberson & CF. Culberson 18171 (holotype:
DUKE!).
(Figs. 51-52)
Holotype description
Thallus pale buff (herbarium), laciniate, loosely adnate, corticicolous, 8 cm broad. Laciniae
dichotomous or irregularly branched, laterally overlapped to somewhat crowded, 1.0-5.5 mm
wide, surface continuous, smooth, opaque to sublustrous, becoming cracked in the center; apical
zone acute or sometimes truncate; margin sinuous, plane, with a strong black line (Fig. 52).
Maculae distinct, effigurate or sometimes forming strong rounded spots, laminai, originating
cracks or not. Cilia black, simple or sometimes furcated, up to 5 mm long, usually thickened,
0.05-0.20 mm wide, frequent. Soralia capitate, orbicular or forming irregular agglomerations,
subapical or laminai; soredia subgranular. Lacinules, pustules and isidia absent. Medulla white.
Undersurface black, lustrous, smooth to rugose, with cracks, widely naked; marginal zone
absent or brown, lustrous, 1-5 mm wide, naked, with a sharp or attenuated limit, smooth to
slightly rugose; rhizines black, irregular in size and ramification, simple to variously branched,
up to 5 mm long, thin or thickened, 0.01-0.30 mm wide, sparse.
Apothecia unknown. Pycnidia submarginal or laminai, rare, ostiole black; conidia not found
(five pycnidia examined).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (minor), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema param ore li en se can be recognized by the laciniate, sorediate thallus,
effigurate maculae (Fig. 52), black and widely naked undersurface, long cilia (up to 5 mm) and
variedly branched, sometimes very thick rhizines (up to 0.3 mm).
This is a strange species inside the group of Parmotrema with salazinic acid, being probably
more related to the genus Everniastrum Hale ex Sipman. Although the effigurate maculae and
dimorphic rhizines could place P. paramoreliense together with the species of the former genus
Rimeliella, the remaining features have no parallel in this group. Future investigations in the
ascospores and conidia, not known up to now, can help to clarify the positioning of this taxon.
Although Culberson & Culberson (1981) asserted that pycnidia are relatively common in this
species, they are rare in the holotype, and no conidia could be seen.
Distribution: Américas (Culberson & Culberson 1981).
64
Parmotrema pectinatum Jungbluth & Marcelli
Bibliotheca Lichenologica 96: 220. 2007.
Type: Brazil, São Paulo State, Municipality of Itirapina, SP-225 Road, Km 111, on a large red
arenite rock emergent above the cerrado forest canopy, in direct sunlight, 22°15'S, 47°49'W, 770
m alt, 16.VI.1979,MP. Marcelli, K. Kalb&A.E. Luchi 16077 (holotype: SP!).
(Fig. 53)
Chemistry: usnic acid, salazinic acid, consalazinic acid and gyrophoric acid (Marcelli et ai.
2007).
This species was recently described for São Paulo State (Marcelli et ai 2007) and is
characterized by a laciniate thallus with narro w laciniate (0.8-3.0 mm), and abundantly ciliate
lobes. It is cio se to P. nylanderi, especially regarding the chemistry and the soredia produced
from pustules. See comments under this species for more details.
Additional specimens examined: Same data as from the holotype, M.P. Marcelli, K. Kalb &
A.E. Luchi 16075, 16079 (SP).
Parmotrema permaculatum (Hale) Kurok.
Bulletin of the National Science Museum Tokyo, Ser. B, 27 (1): 8. 2001.
Parmelia permaculata Hale, Phytologia 21 (6): 425. 1971. - Type: México, State of Veracruz, 9
Km East of Jalapa along highway 140, 1240 m alt., open pasture, scattered cactus and Acácia, on
deciduous trees, 13. III. 1960, leg. M.E. Hale & T.R. Soderstrom 19406 (holotype: US!; isotypes:
DUKE!, S,TNS, UPS).
(Fig. 54)
Holotype description
Thallus milky brown, lobate, loosely adnate, corticicolous, 15 cm broad. Lobes irregularly
branched, laterally overlapped, 4-17 mm wide, surface continuous, smooth or with tiny
depressions, lustrous, becoming rugose and reticulately cracked towards the center, sometimes
with flaking off areolae; apical zone rounded; margin sinuous to broadly crenate, sometimes with
deep incisions, undulated. Maculae distinct, effigurate, laminai, originating cracks. Cilia black,
sometimes pruinose, simple, up to 2 mm long, 0.04-0.10 mm wide, abundant. Lacinules,
pustules, soredia and isidia absent. Medulla white. Undersurface black, opaque to lustrous
(satin-like), smooth, rugulose or papillate, with cracks; marginal zone brown, sometimes white
mottled, lustrous, 3-7 mm wide, naked, with attenuated limit, smooth to rugulose or minutely
veined; rhizines concolor to the undersurface or sometimes with whitish ápices, simple to
irregularly branched, up to 1.5 mm long, 0.01-0.10 mm wide, abundant, evenly distributed.
Apothecia cupuliform, up to 15 mm in diameter, stipitate, laminai to submarginal, margin
slightly incised to lacerate, usually involute, amphithecium maculate, smooth to scrobiculate or
longitudinally rugose and foveolate; ascospores ellipsoid, 13-17 x 7-9 um, episporium 0.5-
1.0 um. Pycnidia submarginal, conspicuous, without prominent margin, frequent, ostiole black;
conidia filiform, 10-15 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
65
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), norstictic acid (trace).
Remarks: Parmotrema permaculatum is distinguished by the effigurate maculae, simple cilia,
black undersurface and absence of vegetative propagules.
It is morphologicaUy similar to P. expansum, which also develops effigurate maculae but has
furcated cilia and shorter conidia (7-12 um).
To compare with another species usually confused in the literature, see comments under P.
enrysacum.
The isotype of P. permaculatum in DUKE was not listed in the protologue (Hale 1971). It is
similar in every respect with the holotype, being clearly conspecific, but develops more abundant
pycnidia and the venation in the undersurface is more evident.
Distribution: United States and México (Hale 1971, Kurokawa 2001).
Parmotrema petropoliense ( Zahlbr.) Spielmann & Marcelli, comb. nov.
Parmelia petropoliensis Zahlbr., Sitzungsberichte der Kaiserlichen Akademie der
Wissenschaften Wien, Mathematisch-Naturwissenschaftliche 111: 426. 1902. - Type: Brazil,
Rio de Janeiro State, Fazenda Inglês prope [Municipality of] Petrópolis, in forest, corticicolous,
leg. Hõhnel 111 (holotype: W!).
(Figs. 55-56)
Holotype description
Thallus pale brown to brown (herbarium), lobate, loosely adnate, corticicolous, 5-7 cm broad.
Lobes irregularly branched, laterally overlapped to quite crowded, 3-6 mm wide, surface
continuous or irregularly cracked, smooth or with tiny depressions, lustrous, becoming
irregularly rugose and reticulately cracked towards the center; apical zone rounded; margin
crenate to incised or lacinulate, undulated. Maculae usually weak, distinct in some lobes,
effigurate, laminai, originating cracks or not. Cilia black, simple or frequently furcated, up to 3
mm long, 0.02-0.10 mm wide, abundant. Lacinules usually simple or irregularly branched,
plane, apex rounded or ± truncate, 1.0-4.0 x 0.5-3.0 mm. Soredia originated from pustuloid
raised structures, marginal to subapical, rarely laminai; soredia granular, commonly developing
into granules, frequently united in agglomerations and forming cilia. Isidia absent. Medulla
white. Undersurface pale brown or dark reddish brown, lustrous, smooth, papillate or slightly
rugose in some áreas, without cracks; marginal zone absent or usually darker than the center;
rhizines black, dimorphic, the shorter ones simple or furcated to dichotomous, usually curled, up
to 1 mm long, thin, 0.01-0.04 mm wide, abundant, evenly distributed; the longer ones up to 1.5
mm long, thickened, 0.05-0.10 mm wide, simple or irregularly branched, few, distributed in
groups.
Apothecia absent. Pycnidia submarginal, inconspicuous, rare, ostiole black; conidia not
found.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
66
Remarks: Parmotrema petropoliense is recognized by the marginal to submarginal pustuloid
raised structures that give rise to soredia, long (up to 3 mm), abundant and frequently furcated
cilia, the brown undersurface with dichotomous rhizines and the effigurate maculae.
This species has long been recognized as a synonym of P. subsumptum, which has extensive
soralia growing in the undersurface. For more details and to differentiate also from P.
margaritatum, see the comments underP. subsumptum.
The holotype collection is constituted by three thalli, glued in a paper card. They are well-
preserved and developed, although without apothecia, and one thallus is glued by the upper
surface.
Distribution: Brazil, Rio de Janeiro State (Zahlbruckner 1902).
Parmotrema pontagrossense (Eliasaro & Adler) Blanco, Crespo, Divakar, Elix & Lumbsch
Mycologia97(l): 157.2005.
Rimelia pontagrossensis Eliasaro & Adler, Mycotaxon 66: 127. 1998. - Type: Brazil, Paraná
State, Municipality of Ponta Grossa, Buraco do Padre, corticicolous, 25°07'S, 50°15'W, 27
January 1996, leg. S. Eliasaro & M. Adler 1621 (holotype: UPCB-28650!; isotypes: BAFC-
37879!, CANB).
(Fig. 57)
Holotype description
Thallus stramineous (herbarium), lobate, more or less closely adnate, corticicolous, 11x7 cm
broad. Lobes irregularly branched, laterally overlapped to somewhat overlapping, 5-13 mm
wide, surface continuous or cracked, smooth to rugose and scrobiculate, lustrous, becoming
almost completely reticulately cracked in the center; apical zone usually rounded; margin crenate
to sublacinulate, slightly plane to undulated. Maculae distinct to strong, punctiform to effigurate,
laminai, sometimes covering almost ali the surface, usually connecting in pseudo-reticular
formations. Cilia black, simple or more frequently 2-3-furcated or cespitose, 0.2-1.5 x 0.02-
0.05 mm, abundant. Isidia, lacinules and pustules absent. Soralia marginal, usually interrupted,
sometimes extensive, rarely orbicular; soredia granular. Medulla white. Undersurface black,
lustrous, rugose or papillate, with some cracks; marginal zone dark brown, 1-3 mm wide, naked,
rarely papillate or with small rhizines, with an attenuated limit; rhizines black, simple, 0.5-2 (-4)
x 0.01-0.1 mm, abundant, apparently evenly distributed, but very dense near the marginal zone.
Apothecia unknown. Pycnidia submarginal, little conspicuous, without prominent margin,
few, ostiole black; conidia not found (ca. 20 pycnidia investigated; the isotype from BAFC! has
filiform conidia, 9-11 x ca. 1 .0 um).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV+ yellow.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), lichexanthone (minor), lichesterinic acid (major), protolichesterinic acid (major).
Remarks: Parmotrema pontagrossense is distinguished by the sorediate thallus with cespitose
cilia and by presents a complex chemistry, with salazinic acid, lichexanthone (see comments
underP. ultralucens) and the lichesterinic and protolichesterinic acids in the medulla.
The branching pattern of the cilia was already well described in the protologue (Eliasaro &
Adler 1998). The same pattern can be seen in Parmotrema spinibarbe, a close species with the
67
same overall morphology, but that can be differentiated by the formation of soredia from
botryose structures (Fig. 71) as well as by the absence of lichesterinic and protolichesterinic
acids. Future collections of fertile material perhaps will provide additional differences between
the se two species.
The types studied are well developed and completely similar. The undersurface is difficult to
access since the thalli are closely adnate to the substrate; however, the free zone cio se to the
margins presented sufficient diagnostic features.
Distribution: known only for Brazil, where it was recorded to the States of Paraná (Eliasaro &
Adler 1998, Donha 2005) and Rio Grande do Sul (Canêz 2005).
Parmotrema pseudoreticulatum (Tavares) Hale
Phytologia28 (4): 338. 1974.
Parmelia pseiidoreticiãata Tavares, Portugaliae Acta Biológica, série B, 1 (1-2): 138. 1945. -
Type: Portugal, surroundings of Lisboa, leg. C. Tavares (holotype: LSU, syntype, F!).
= Parmelia perlata var. olivaria A eh., Meth. Lich.: 217. 1803. Nom. rej. prop. (Hawksworth et
ai. 2003). - Type: Spain, sine loc, on Olea, 1971-93, leg. P.K. A. Schousboê (lectotype,
selected by Hawksworth et ai 2003: H-ACH 1327c), fide Divakar et ai (2005).
= Parmelia olivaria (Ach.) Hue, Nouv. Arch. Mus. Hist. Nat. Paris, sér. 4 (1): 195. 1899. Fide
Divakar et ai (2005).
= Rimelia olivaria (Ach.) Hale & Fletcher, The Bryologist 93 (1): 28. 1990. Fide Divakar et ai
(2005).
(Fig. 58)
Syntype description (F)
Thallus buff (herbarium), lobate, apparently loosely adnate, corticicolous, 8.5-7.0 cm broad.
Lobes irregularly branched, laterally overlapped to somewhat crowded, 4-12 mm wide, surface
continuous, smooth or irregular, lustrous, becoming rugose and cracked in the center; apical zone
rounded, usually concave; margin smooth to crenate or broadly crenate, sometimes deeply
incised, undulated. Maculae distinct, punctiform or effigurate, sometimes assuming a reticular
pattern, laminai, in some áreas covering almost ali the surface (hypermaculate). Cilia black,
simple, up to 0.5 mm long, rare. Sor alia marginal and linear or more commonly laminai and
capitate, sometimes extensive; soredia subgranular. Lacinules, pustules and isidia absent.
Medulla white. Undersurface black, lustrous, usually finely rugose, with few cracks; marginal
zone pale brown, lustrous, 1-5 mm wide, naked or rarely rhizinate, with an attenuated limit,
smooth, rugose or papillate; rhizines black, simple, up to 1 mm long, abundant, evenly
distributed.
Apothecia absent. Pycnidia submarginal, conspicuous, without prominent margin, abundant,
ostiole black; conidia filiform, 8-14 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), protocetraric acid (trace).
68
Remarks: P armo trema pseiidoreticidatiim is distinguished by the abundant laminai and capitate
soralia, the rare short cilia (up to 0.5 mm), the punctiform to effigurate maculae and the naked
under marginal zone.
At first sight, this seems to be a typical Rime lia; however, it is easy to verify that the (false)
reticulation is formed by the confluence of the maculae, i.e., the maculae are actually punctiform
or effigurate as those of Canomaculina.
P armo trema pseiidoreticidatum was considered to be a synonym of P. reticulatum by Krog &
Swinscow (1981), which has distinct reticulate maculae, under marginal zone rhizinate, soralia
growing in short lacinules and more frequent cilia. The distinction between them was also
molecularly supported (Hawksworth et ai. 2003, Divakar et ai. 2005).
The syntype here described was not listed in the pertinent literature. It is well-preserved and
developed, although sterile, and glued on a paper card.
Distribution: Europe and Azores (Tavares 1945, Hawksworth et ai. 2003) and Africa (Divakar
et ai. 2005).
Parmotrema radiatum (Lynge) Spielmann & Marcelli, comb. et stat. nov.
Parmelia cetrata Ach. subsp. radiata Lynge, Arkiv for Botanik 13 (13): 94. 1914. - Type:
Paraguay, Colónia Risso prope Rio Apa, corticicolous, 23. IX. 1893, leg. G.O.A. Malme 1834
(lectotype: S!; duplicate of the lectotype: UPS!).
(Fig. 59)
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, corticicolous, 10.5 cm broad. Lobes
irregularly branched, laterally overlapped, distinctly radial, 3.0-7.5 mm wide, surface
continuous, smooth, lustrous, frequently pruinose in the lobe ápices, becoming slightly rugose
and reticulately cracked towards the center; apical zone rounded; margin crenate to sinous-
incised, undulated. Maculae distinct, reticular to extensive (hypermaculate), laminai, marginal,
originating cracks. Cilia black, usually squarrose, less frequently simple or furcated, up to 1.5 (-
2.0) mm long, 0.03-0.10 mm wide, abundant. Lacinules, pustules, soredia and isidia absent.
Medulla white. Undersurface black, lustrous (satin-like), smooth, sometimes papillate or veined,
with abundant cracks; marginal zone dark brown, shiny, narro w, 1-2 mm wide, naked or with
sparse rhizines, with attenuated limit, smooth or papillate; rhizines black, usually strongly
squarrose, sometimes irregularly branched or simple, up to 3 mm long, thin to thickened, 0.01-
0.15 mm wide , abundant, irregularly distributed.
Apothecia cupuliform, 1-8 mm in diameter, stipitate, submarginal, margin smooth to dentate,
amphithecium maculate, smooth to rugose, disc brown, epruinose, perforate at maturity;
ascospores ellipsoid, 11.0-13 x 7.5-9.0 um, episporium 1.0-1.5 um. Pycnidia submarginal,
rarely laminai, conspicuous, usually with a prominent margin, frequent, ostiole black; conidia
filiform, 10-15 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), protocetraric acid (trace).
69
Remarks: the more salient features of Parmotrema radiatum are the usually squarrose cilia, the
upper surface reticulate-maculate or hypermaculate, the squarrose and richly branched rhizinae
and the shiny nude dark brown under margin.
This species was included by Hale & Fletcher in the synonymy of P. cetratum. Nevertheless,
P. cetratum presents only reticular maculae, cilia never squarrose and the marginal zone is
totally absent (lectotype: H-ACH).
Lynge (1914) described Parmelia cetrata subsp. radiata based in one specimen from Brazil
and another from Paraguay. The Brazilian taxon belongs to a new species, here described (see P.
bifidum).
The duplicate of the lectotype agrees very well with the lectotype, including ascospores and
conidial data. It is clearly a part of the holotype thallus. However, the chemical analysis could
not demonstrate the presence of trace of protocetraric acid.
Distribution: Paraguay (Lynge 1914).
Parmotrema ramusculum (Hale) Hale
Phytologia 28 (4): 338. 1974.
Parmelia ramuscula Hale, Contributions from the United States National Herbarium 36 (5): 261.
1965. - Type: Borneo Island, Malaysia, Sarawak state, Lawas, on a Rubber tree. 31.V.1955, leg.
W.M.A Brooke 10031 (holotype: BM!; isotype: L!, US).
(Fig. 60)
Holotype description
Thallus grayish, lobate, loosely adnate, corticicolous, 9.5 cm broad. Lobes irregularly branched,
4-10 mm wide, surface continuous, smooth, sublustrous, becoming irregularly cracked towards
the center; apical zone rounded; margin smooth. Maculae weak, punctiform, laminai, more
easily seen in young lobes. Cilia black, simple, up to 1.3 mm long, 0.04-0.10 mm wide, sparse.
Lacinules present, forming arbuscular, sorediate and pustular structures, irregularly branched,
usually terete or sometimes ílattened, up to 1.5 cm long, individual branches 0.2-1.0 mm wide.
Soredia whitish, farino se or becoming more compact, developed in the arbuscular structures and
in some lobe margins. True isidia absent. Medulla white. Undersurface (based on the L isotype):
black, lustrous, smooth to slightly rugose or rarely papillate, with numerous cracks; marginal
zone absent (black) or eburneus under the lobes that give rise to the arbuscular structures, which
are also usually eburneus; rhizines black, simple, up to 1 x 0.05 mm, rare, distributed in groups.
Apothecia and pycnidia unknown.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: P ramusculum is recognized by the very distinct arbuscular structures that became
entirely sorediate.
Although these structures were already named as "coralloid isidia" (Hale 1965), they
probably deserves an own name, "arbuscula", since are different from anything that could be
called "isidia" in another species. Similar structures are found in P. coralliforme, although in this
case they are usually ciliate and not so sorediate as in P. ramusculum. Additionally
70
P. coralliforme has distinct effigurate maculae, abundant cilia and rhizines and a pale brown
marginal zone in the undersurface.
The two other comparable species are P. flavotinctum (Hale) Hale (with atranorin only) and
P. fasciciãatum (Vainio) Hale (with a very complex chemistry, without salazinic acid, see
Louwhoff & Elix 1999). They need also to be reevaluated before these intriguing structures
could be adequately named.
A well-developed and preserved specimen represents the isotype (L). It has subcanaliculate
lobes and more commonly furcated cilia, but in the other features is identical to the holotype.
Since it is not glued as the holotype, it was used to describe the undersurface.
Distribution: Malaysian Borneo and Philippines (Hale 1965) and Guyana (Sipman 1999).
Parmotrema reitzii Hale
Mycotaxon 5 (2): 439. 1977.
Type: Brazil, Santa Catarina, [Municipality of] Lajes, Locality of Encruzilhada, 950 m alt., on
bark, 24.IV.1962, leg. R. Reitz & R.M. Klein 12927 (holotype: US!).
= Canomacidina reitzii (Hale) Elix, Mycotaxon 65: 477. 1997.
= Rimeliella reitzii (Hale) Fletcher, in DePriest & B. Hale, Mycotaxon 67: 205. 1998.
(Figs. 61-62)
Holotype description
Thallus pale brown (herbarium), lobate, loosely adnate, corticicolous, 12-14 cm broad. Lobes
irregularly branched, laterally overlapped , 0.4-17 mm wide, surface continuous, smooth or with
± rounded depressions, lustrous, becoming reticulately cracked and sometimes rugose towards
the center; apical zone rounded; margin crenate-incised to sublacinulate, undulated. Maculae
distinct, effigurate, laminai, originating cracks. Cilia black, simple or rarely with a lateral
branchlet, up to 1.0 (-1.5) mm long, thin to somewhat thickened, 0.01-0.10 mm wide, frequent
to few. Soralia whitish, marginal, the soredia usually pilling up to form raised arbuscular or
richly branched, partially corticated structures, turning the lobe ápices involute, assuming an
linear pattern when coníluent; soredia subfarinose. Lacinules, pustules and isidia absent.
Medulla white. Undersurface pale brown (perhaps white in fresh material), sublustrous, smooth
to papillate, without cracks; marginal zone absent; rhizines black or rarely dark brown,
dimorphic, 1) the shorter ones simple or sometimes furcated, usually curled, up to 0.5 mm long,
thin, 0.01-0.04 mm wide, abundant, evenly distributed; 2) the longer ones up to 3.5 mm long,
thickened, 0.05-0. 20 mm wide, simple or more commonly irregularly branched, especially near
the apex, frequent, distributed in groups.
Apothecia and pycnidia unkno wn.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloro atranorin (trace), salazinic acid (major), consalazinic acid
(minor), norlobaridone (minor), loxodin (trace).
Remarks: Parmotrema reitzii is recognized by the submarginal soralia, usually raised and
forming branched structures, the pale brown to brown underside, with dimorphic rhizines, and
the medullar chemistry, with salazinic acid and norlobaridone.
71
Parmotrema petropoliense produces similar sorediate structures but with different ontogeny,
and has distinct lacinules and furcated cilia, besides the absence of norlobaridone.
Another cio se species is P. subsumptum (see comments and table under this species), where
the soralia grow in the underside of the crisped lobes, and additionally do not produces
norlobaridone.
Kurokawa (1991) placedP. reitzii as synonym of Parmotrema conferendum Hale, while Fleig
(1997) situated it as synonym of P. subsumptum. The type of P. conferendum was not studied
here, but with base in the type of P. subsumptum and its synonyms, P. reitzii is distinct booth
morphologically (especially the soralia ontogeny) and chemically, i.e., by the production of
norlobaridone.
Distribution: South America, where it was recorded to Brazil (Hale 1977, Marcelli 2008) and
Venezuela (Feuerer 2008). In Brazil it is known to the States of Rio Grande do Sul (Fleig 1997),
Santa Catarina (Hale 1977).
Parmotrema reparatum (Stirton) Blanco, Crespo, Divakar, Elix & Lumbsch
Mycologia97(l): 157.2005.
Parmelia reparata Stirton, Scottish Naturalist 4: 201. 1877-78. - Type: Austrália, Queensland,
Cave Mountain near Brisbane, leg. F.M. Bailey 16 (lectotype: BM!).
= Canomaculina reparata (Stirton) Kurok., Bulletin of the National Science Museum Tokyo,
Ser. B, 27(1): 2. 2001.
= Parmelia virens Míill. Arg., Flora 69: 255. 1886. - Type: Austrália, Queensland, Toowoomba,
1882, leg. Hartmann (holotype: G).
(Fig. 63)
Lectotype description
Thallus brown (herbarium), lobate, loosely adnate, corticicolous, 7 cm broad. Lobes irregularly
branched, laterally overlapped, 3-9 mm wide, surface continuous, smooth, lustrous, becoming
scrobiculate and irregularly cracked towards the center; apical zone rounded; margin broadly
crenate, undulated. Maculae distinct, effigurate, laminai. Cilia black, simple or rarely furcated,
up to 1.5 mm long, 0.04-0.07 mm wide, frequent. Lacinules, pustules, soredia and isidia absent.
Medulla white. Undersurface black, opaque, extensively veined, without cracks; marginal zone
absent (i.e., black) or dark brown to brown, then opaque to lustrous, 3-7 mm wide, rhizinate, or
papillate; rhizines black or dark brown in the marginal zone, dimorphic, the short ones simple
and usually curled, up to 0.5 mm long, 0.01-0.04 mm wide, abundant and evenly distributed, the
longer ones simple or branched, up to 1.5 mm long, 0.04-0.10 mm, frequent, distributed in
groups.
Apothecia cupuliform, 2-13 mm in diameter, stipitate, submarginal, margin smooth to slightly
incised or lacerate, usually involute, amphithecium maculate, scrobiculate to longitudinally
rugose and with deep depressions, disc brown, epruinose, perforate; ascospores subglobose,
10.0-12.5 x 8.0-9.0 um, episporium ca. 1 um. Pycnidia mainly submarginal, conspicuous, with
or without prominent margin, frequent; conidia filiform, 9-15 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
72
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema reparatum is recognized by the effigurate maculae, simple cilia, black
undersurface, dimorphic rhizines and the absence of vegetative propagules.
Parmotrema subcaperatum is quite similar in overall morphology, but the marginal zone is
more distinct, the ascospores larger (14-16 um), and additionally contains usnic acid.
Parmelia virens has the same overall morphology of P. reparata, except by the darker,
olivaceous upper surface. Therefore, we are accepting the synonymization proposed by
Kurokawa (2001), although some comments are necessary:
When describing Parmelia reparata, Stirton (1877-1878) noted ascospores 13-14 x 9-11 um.
The study of the lectotype revealed ascospores briefly shorter and subglobose, 10.0-12.5 x 8.0-
9.0 um, while the holotype of P. virens presented still smaller measurements, of also subglobose
ascospores, 7.5-12 x 7.0-8.5 um.
The ascospores size published by Kurokawa (2001), however, are quite distant from these:
15-18 x 6-8 um. This clearly represents ellipsoid or more probably narro wly ellipsoid
ascospores. Apparently this measurement should be referred to the specimens studied.
Unfortunately, full comparisons of the undersurface of P. reparata and P. virens are not
possible, since the lectotype of the first species is almost totally glued to a card.
Kurokawa (200 1 ) postulated that P. subcaperatum could be endemic to South America, while
P. reparatum occurs in Austrália and New Zealand. Nevertheless, the specimens at the
"halfway", namely Africa (especially that recorded by Krog & Swinscow 1981), still need to be
revised to corroborate this hypothesis.
The specimens listed in Hale (1965) to the United States of America and México actually
belong to P. permaculatum, as already indicated by Hale (1971).The specimens recorded as
Canomacalina aff reparata by Spielmann (2005) need to be checked, since they have reticular
maculae.
Distribution: Oceania (Kurokawa 2001)
Parmotrema reterimulosum (Zahlbr.) Spielmann & Marcelli, comb. nov.
Parmelia reterimulosa Zahlbr., Botanische Jahrbucher fur Systematik, Pílanzengeschichte um
Pílanzengeographie 60: 520. 1926. - Type: South Africa, Natal, Drackenberge, Van Reenens
Pass, leg. Bnmnthaler s.n. (holotype: WU!).
(Fig. 64)
Holotype description
Thallus brownish (herbarium), lobate, apparently loosely adnate, corticicolous, 7 cm broad.
Lobes irregularly branched, laterally overlapped to crowded, 2-5 mm wide, surface continuous
or irregularly cracked, smooth or slightly irregular, lustrous, becoming strongly cracked in the
center; apical zone rounded; margin sinuous, undulated. Lacinules absent, although the thallus
subdivision could be interpreted as forming lacinules. Maculae distinct, reticular, laminai,
originating cracks. Cilia black, simple, up to 1.5 mm long, abundant, at times some "cilia"
(rhizines?) growing in the underside of the lobes. Pustules absent. Soralia whitish, extensive,
submarginal and moving on to the center, present also in the ápices of the sublacinules; soredia
granular. Isidia absent. Medulla white. Undersurface black, lustrous, rugose or papillate, with
scars; marginal zone brown, or dark-brown to black in some áreas, lustrous, 1-3 mm wide,
naked or with papillate rhizines, with an attenuated limit, smooth to slightly rugose; rhizines
73
black, usually simple, rarely with lateral branches, up to 1.5 mm long, more or less evenly
distributed, in some áreas the rhizines are strongly curly and entangled e sometimes forming
groups of rhizines slightly thickened.
Apothecia pressed down, apparently concave, 1 mm in diameter, substipitate, laminai, margin
smooth crenate, amphithecium strongly rugose, disc dark brown, epruinose, perforate;
ascospores ellipsoid or oblong, 12.5-17.5 x 6.0-7.5 um, episporium ca. 1.0 um. Pycnidia
laminai to submarginal, conspicuous, without prominent margin, frequent, ostiole black; conidia
filiform, 11-16 (-18) x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema reterimiãosiim is recognized by the soredia ontogeny, the reticular
maculae, the bare marginal zone and the filiform relatively long conidia (11-18 um).
This species was proposed by Zahlbruckner (1926), who made important observations and
provided a very complete description. Dodge (1959) accepted it, citing several additional
specimens. Hale & Fletcher (1990) placed it as synonym of P. reticulatum.
However, P. reticulatum has a rhizinate under marginal zone, smaller conidia 8.5-12.0 um
long, and the soredia arise from swellings which become defragmented.
In P. reterimulosum the, córtex become fragmented into minute areolae, which afterwards
give rise to the soredia in extensive soralia. Usually, each areolae give rise to only one soredium.
The presence of the soralia apparently turns down the lobe ápices. The underside marginal zone
is papillate or with very rudimentary immature rhizines, or more commonly naked.
The holotype is constituted by a broken thallus (five parts), and a small envelope with one
apothecium. Together there is an isidiate, ciliate lichen, probably with stictic acid (medulla K+
yellow).
Distribution: Africa (Zahlbruckner 1926, Dodge 1959).
Parmotrema reticulatum (Taylor) M. Choisy
Bulletin mensuel de la Société Linnéenne de Lyon 21 : 175. 1952.
Parmelia reticulata Taylor, in Mackay, J.T., Flora hibernica, part second, comprising the Musci,
Hepaticae and Lichenes, 1836, p. 148. - Type: Ireland, County of Kerry, near Dunkerron, on
rocks, common, leg. T Taylor s.n. (lectotype: FH!, duplicate of the lectotype: BM!).
= Rimelia reticulata (Taylor) Hale & Fletcher, The Bryologist 93 (1): 28. 1990.
= Parmelia praeperlata Nyl., Flora 69: 319. 1886. Type: - French Southern and Antarctic Lands,
St. Paul Island, leg. Fenzl s.n. (lectotype: H-NYL 35548!).
(Fig. 65)
Duplicate of the lectotype description (BM)
Thallus slightly brownish, lobate, loosely adnate, over rock with mosses, 6.5-7.0 x 4.0-5.0 cm
broad. Lobes irregularly branched, crowded, 2.0-8.0 mm wide, surface continuous or reticulately
cracked, smooth, lustrous, usually cracked in the center; apical zone usually subdivided in
74
lacinules, when entire rounded; margin smooth, broadly crenate, undulated. Maculae distinct,
reticular, laminai, originating cracks. Lacinules simple or bifurcate, present in the whole thallus,
plane to slightly canaliculated or rarely concave, usually with the apex involute, often covered by
soredia, apex acute or sometimes rounded, 0.5-2.2 x 0.6-1.6 mm. Cilia black, usually simple,
rare irregularly branched, squarrose or bifurcated in the apex, 0.1-1.0 x 0.02-0.05 (-0.10) mm,
few to frequent in the non-sorediate lobes, absent in the sorediate lacinules. Soralia subapical in
the short lacinules, growing backwards some millimeters in the lamina, becoming labriform in
several áreas; soredia subgranular to farino se, rising from swellings which become
defragmented. Pustides and isidia absent. Medulla white. Undersurface black, lustrous, rugose to
veined, with scars; marginal zone brown or rarely black, lustrous, 0.5-3.0 mm wide, rhizinate,
rugose or papillate, with an attenuated limit; rhizines black, of two extremes: (1) simple to
squarrose, slender (0.02-0.05 mm wide) and usually shorter (0.2-1.3 mm), abundant, evenly
distributed, or (2) irregularly branched, especially close to the ápices, thickened (0,10-0.25 mm)
and longer (0.5-2.0 mm), frequent, distributed in groups.
Apothecia immature (asei and ascospores not formed), infrequent (four found), urceolate, 3.5
mm in diameter, substipitate, laminai, margin dentate, sorediate or in process of soredia
formation, amphithecium sorediate, disc beige, epruinose, imperforate. Pycnidia laminai to
submarginal, conspicuous, without prominent margin, abundant, ostiole black; conidia filiform,
8.5-12.0 x C a. 1.0 fim.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema reticulatum is characterized by reticular maculae, the presence of
submarginal, extensive soralia (which also can oceur in the short lacinules) and the produetion of
salazinic acid.
It is part of a complex of species earlier grouped under Rimelia reticulata. Some very
divergent entities were part of this complex, like pustulate species with reticular maculae (P.
granulare), sorediate species with effigurate maculae (P. concors, P. livido-tesselatum) and
pustulate species with effigurate maculae (P. nudurri).
Also sometimes included under P. reticulatum, P. clavuliferum is distinct by the formation of
closed lacinules, sorediate only at the ápices.
The rearrangement of the taxa involved lead to the conclusion that the present data on the
distribution of P. reticulatum cannot be take into account, and probably this species has a much
more restrict distribution, maybe only in temperate áreas of Europe and North America.
Interestingly the apothecia seen in the types are imperforate. According to Hale & Fletcher
(1990), species of Rimelia would always have perforate apothecia. Taylor (1967) also found only
imperforate apothecia in Rimelia reticulata. However, Purvis et ai. (1992) asserted that they can
be sometimes perforate.
The duplicate of the lectotype of P. reticulatum is composed of two thalli glued in a paper
box. They are similar in soralia, branching, maculation, and conidia size and shape. There are
four immature apothecia. Hale & Fletcher (1990) asserted that the ascospores of Rimelia
reticulata are 13-18 x 8-10 um. However, it is not known from what specimen or specimens
these measures were obtained. The conidia found in the duplicate of the lectotype are shorter
(8.5-12.0 um) than those cited by Hale & Fletcher (1990): 12-16 um. The soralia in the
duplicate of the lectotype reminds Parmotrema simulans (Hale) Hale, which have only atranorin
and caperatic acid and was once thought to be a chemotype of P. reticulatum (Hale & Fletcher
1990).
75
In the free (not glued) undersurfaces, some thickened rhizines were found, branched and
longer in relation to the more common ones. However, a gradual transition between one type to
another can be observed.
The lectotype of P. reticulatum (FH) is formed by three pieces glued in a sheet. Although
agreeing well with the duplicate of the lectotype, it has structures that can resemble schizidia,
since they are areolae that can give rise to soredia at their margins. A re-study is necessary to be
sure about these data. Also, since it does not have apothecia or pycnidia, we chose to present
here the description of the duplicate of the lectotype.
Parmelia praeperlata Nyl., known only from the type locality, was published by Nylander
(1875), and a more complete description was presented some years later (Nylander 1886). It is
interesting to note that Hale & Fletcher (1990) indicated the paper of Nylander (1886) as the
protologue, not Nylander (1875). It was the Catalogas (Zahlbruckner 1930a) that called our
attention to these papers.
The lectotype of P. praeperlata has a strong black, marginal line, and sometimes develops
short, branched tapered cilia (up to 0.5 mm), which are a noticeable feature. However, P.
reticulatum also develops a dark brown to black marginal line and similar branched cilia,
although just in some lobes. A label by J.A. Elix, dated November 09, 1998, gives the following
chemical data: "TLC solvent C/HPLC: atranorin (minor), chloroatranorin (minor), salazinic acid
(major), consalazinic acid (minor)". Nylander (1886) reported the conidia of Parmelia
praeperlata as 9-11 um. We found it as 8.5-12 x ca. 1 um. So the same size asP. reticulatum.
Since apothecia are lacking or immature in the types of P. praeperlata and P. reticulatum,
additional date could not be compared and so it seems better to keep these species as synonyms,
until more specimens be available.
Parmotrema ruminatum (Zahlbr.) Blanco, Crespo, Divakar, Elix & Lumbsch
Mycologia97(l): 158.2005.
Parmelia ruminata Zahlbr. in Magnusson & Zahlbruckner, Arkiv for botanik 3 IA (6): 107.
1944. - Type: Hawaiian Islands, 200 feet alt. (656 m), leg. Brother Matthias 757 (lectotype: W!,
selected here).
= Rimelia ruminata (Zahlbr.) Hale & Fletcher, The Bryologist 93 (1): 28. 1990.
= Parmelia ramescens Zahlbr. in Magnusson & Zahlbruckner, Arkiv for Botanik 31 A (6): 97. -
Type: Hawaii, Maui, Iao Valley, leg. Faurie 516 (lectotype: W!; duplicates of the lectotype:
BM!,PC).
= Rimelia ramescens (Zahlbr.) K.H. Moon & Kurok., Journal of Japanese Botany 76: 326.
(Fig. 66)
Lectotype description
Thallus brownish (herbarium), lo bate, 10.5 cm broad. Lobes irregularly branched, ± crowded, 2-
5 mm wide, surface continuous to cracked, slightly rugose or irregular, becoming strongly
rugose, cracked and pustulate towards the center, sometimes producing areolae that ílake o ff;
apical zone rounded, usually pruinose; margin smooth to sinuous or sublobulate, undulated.
Maculae distinct, reticular, laminai, originating cracks. Cilia absent to rare (see comments
below). P ustules rugose, capitate or elongated, laminai, remaining entire or sometimes flaking
off together with the areolae. Lacinules, soralia and isidia absent. Medulla white. Undersurface
black, lustrous, smooth to rugose, with cracks; marginal zone brown, lustrous, 1-2 mm wide,
76
naked, with an attenuated limit, smooth; rhizines black, simple, up to 1 mm long, 0.01-0.04 mm
wide.
Apothecia cupuliform, \-A mm in diameter, short-stipitate, laminai, margin crenate, pustulate,
amphithecium rugose, pustulate, disc brown, epruinose, imperforate; ascospores oblong-
ellipsoid, 16-20 x 7.5-9.0 um, episporium 1.0-1.5 um. Pycnidia submarginal, conspicuous,
usually without prominent margin, few, ostiole black; conidia filiform, 9-13 x ca. 1 .0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC (J. Johnston & G.A. Jenkins, 03.11.1988, label with the lectotype in W):
atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (trace), protocetraric acid
(trace), unknown RT 5.81 (trace, HPLC only).
Remarks: Parmotrema ruminatum is characterized by the strongly pustulate upper surface, from
which sometimes areolae are ílaked off, and the eciliate lobe margins.
It is related to P. austrocetratum, in which the production of areolae is much more evident,
giving rise to schizidia, and the cilia are abundant and longer (up to 1.6 mm).
The lectotype of P. ruminatum is well developed and shows ali features necessary to its
description, with exception of the undersurface, since the thallus is strongly glued to a paper
card. Although a greenish fungus is abundant in the upper surface, it seems it is not causing any
damage to the thallus. Rare black, simple cilia, 0.1-0.6 x 0.02-0.05 mm were found. These look
much more as an "accident" (some rhizinate growing in the margin) that true cilia, and so the
species is considered eciliate here.
The ascospores of this species, oblong-ellipsoid, are quite uncommon in the Parmotrema with
salazinic acid. So strange that we first thought the measurements described in Magnusson &
Zahlbruckner (1944) could be wrong, but confirmation through the lectotype study then
confirmed that data.
The typification revealed some problems. Magnusson & Zahlbruckner (1944) listed two
collections, one from Hawaiian Islands (Brother Matthias) and one from Hawaii (Faurie no.
861). According to a label from Hale (dated 1965), the specimen Brother Matthias 757 (W) is
the holotype. However, Hale & Fletcher (1990) chose as lectotype "Brother Matthias (Faurie
861), Hawaii". We do not know where the specimen Faurie 861 is preserved, and future studies
must trace it to clarify it status. Thus, for practical purposes, the specimen Brother Matthias 757
is here being selected as lectotype.
Both the lectotype and duplicate of the lectotype of P arme lia ramescens are well developed
thalli, the specimen from W being 10 cm broad and the BM specimen 13 cm broad. The upper
surface is strongly rugose and pustulate, and some lobes become canaliculated. In the other
features they agree well with the holotype of P. rumina ta, and so are here being treated as its
synonym, a position already proposed by Hale & Fletcher (1990). Zahlbruckner (in Magnusson
& Zahlbruckner 1944) asserted that he saw a large thallus up to 20 cm broad. Maybe this is the
specimen deposited in PC, since we have not received it.
Conidia were found only in the duplicate of the lectotype of P. ramescens (BM). They are
filiform, 9-14 x ca. 1 um, so matching the conidia of P. ruminata.
Moon et ai. (2001) proposed Rimelia ramescens as a distinct species. The authors asserted
that it has "a rather smooth upper surface" and "ií lacks pustules and eminent ridges". This
observation, however, do not agree with ours, made on the lectotype and its duplicate. It is
possible that they refer to the specimen examined only. The herbarium where the lectotype is
hosted is actually W, not TNS, as referred by those authors.
Parmotrema ruminatum has an indeterminate substrate. It was quoted as "saxicolous?" by
Magnusson & Zahlbruckner (1944), corticicolous by Moon et ai. (2001) and finally both in
77
Smith (1993). This last author, nevertheless, asserted that the species is schizidiate, not pustulate,
and so his identification needs to be confirmed.
Distribution: Hawaiian Islands (Magnusson & Zahlbruckner 1944, Smith 1993, Hale & Fletcher
1990, Moon et ai. 2001).
Parmotrema ruptum (Lynge) Hale ex DePriest & B. Hale
Mycotaxon 67: 204. 1998.
Parmelia rupta Lynge, Arkiv for Botanik 13 (13): 40. 1914. - Type: Paraguay, Territory Gran
Chaco, ad Rio Negro, 14.IX.1893, corticicolous, leg. G.O.A. Malme (holotype: S!).
(Fig. 67)
Holotype description
Thallus brownish (herbarium), lobate, ± adnate, corticicolous, 6 cm broad. Lobes irregularly
branched to quite crowded in the center, 2-7 mm wide, surface continuous or reticulately
cracked, smooth to irregular, lustrous, becoming fully reticulately cracked towards the center;
apical zone rounded; margin smooth to sublacinulate or lobulate (see comments), undulated.
Maculae distinct, reticular to hypermaculate, laminai, originating cracks. Cilia black, simple or
rarely furcated near the apex, short, up to 0.5 mm long, usually thin but sometimes thickened,
0.02-0.10 mm wide, few. Sublacinulae or lobules simple, few, plane, apex usually acute to
rounded, rarely truncate, 0.2-2.0 x 0.2-1.0 mm. P ustules, soredia and isidia absent. Medulla
white. Undersurface black, lustrous, rugose, subscrobiculate, papillate or veined, with cracks;
marginal zone dark brown to dark reddish brown, lustrous, 0.5-2.0 mm wide, naked, with an
attenuated limit, smooth to slightly rugose; rhizines black, usually simple, up to 1 mm long, thin
to thickened, 0.02-0.15 mm wide, abundant, evenly distributed.
Apothecia plane to concave or cupuliform, 1.5-5.5 mm in diameter, short- stipitate, usually
submarginal, margin smooth, crenate or deeply incised, dividing the apothecium in parts, rarely
with small lobules, amphithecium maculate, smooth or rugose, disc pale brown, epruinose,
usually rugose at maturity, imperforate although with a central depression; ascospores
subglobose to ellipsoid, biserial, 10.0-12.5 x 6.0-7.5 um, episporium ca. 1 um. Pycnidia
submarginal or laminai, conspicuous, with or without prominent margin, abundant, ostiole black;
conidia filiform, 10.0-12.5 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC (J.A. Elix, 28.XII.2007, label with the holotype): atranorin (minor),
chloroatranorin (minor), salazinic acid (major), consalazinic acid (minor).
Remarks: Parmotrema ruptum can be recognized by the strongly reticulately cracked upper
surface, associated with reticulate maculae (sometimes hypermaculate), the absence of
propagules, the short, rare cilia, the naked under margin, usually dark reddish brown, and the
production of salazinic acid.
Fleig (1997) consideredP. ruptum andP. expansum synonyms of P. eurysacum. However,
according the types studied,P. eurysacum is emaculate, andP. expansum has effigurate maculae.
It also could be confused withP. cetratum, which is distinctly rhizinate up to the margin.
The specimens recorded by Canêz (2005) as Parmotrema ruptum were studied (SP). They are
represented by well-developed thalli. In fact, one thallus is 22 cm broad. The upper surface is
78
usually strongly maculate (hypermaculate) and whitish, resembling P. albinatum, a sorediate
species recently described from Hawaii (Moon et ai. 2001), although in some lobes the reticular
maculae are conspicuous. The cilia are quite frequently long (up to 1 mm) and bifurcate, and the
thallus is almost uniformly whitish. Nevertheless, the underside is practically identical to the
holotype de P. ruptum, especially the dark reddish brown under margin and rugose surface. The
conidia also are equal. So they are conspecific.
It must be noted that ali specimens recorded by Canêz (2005) are saxicolous, while the
holotype of P. ruptum is corticicolous. Unfortunately, the specimens reported by Canêz (2005)
don't developed apothecia, avoiding the comparison of ascospores data.
The holotype of P. ruptum is attached to a bark, so some the undersurface features were not
described in detail, the observations being based only on the free lobes. The thallus also form
some adventitious lobules or sublacinules, rose on the damaged áreas. In future collections of
this species special attention must be paid in these structures (if present) and also on apothecial
margins. They are quite similar to the lobe margins, and maybe it can give rise to apothecial
cilia.
Distribution: Brazil (Canêz 2005), Paraguay (Lynge 1914, Hale 1960), Uruguay (Rásánen 1938,
Osório 1972).
Additional specimens examined: Brazil, Rio Grande do Sul, Municipality of Vacaria, Fazenda
da Estrela, 28°04'49.2"S, 50°56'36.9"W, 850 m, over basaltic rock, sunny place, 10.1.2004, leg.
L.S. Canêz &A.A. Spielmann 1020, 1021, 1035, 1080 (SP).
Parmotrema sieberi (C.W. Dodge) Spielmann & Marcelli, comb. nov.
Parmelia sieberi C.W. Dodge, Annals of the Missouri Botanical Garden 46: 148. 1959. -Type:
Mauritius, leg. Sieber, Crypt. Exot. 44, corticole, ex herb. Sbarbaro (holotype and isotype: FH!).
= Parmelia claudelii (Harm.) Vainio var. clemensiae Vainio, Phil. Journ. Sei. 4: 659. 1909. -
Type: Philippines, Mindanao, Lake Lanao, leg. Clemens 1319 (lectotype: TUR, Vainio herb.
n°2542).
(Fig. 68)
Holotype description
Thallus brown, lo bate, loosely adnate, apparently corticicolous, 45 cm broad. Lobes irregularly
branched, laterally overlapped, 8-25 mm wide, surface continuous or less frequently cracked,
usually smooth, lustrous, becoming rugose and cracked in the center; apical zone rounded;
margin smooth or sparsely crenate, undulated. Cilia black, simple or furcated, sometimes more
branched, up to 2 mm long, frequent, less dense in the lobe ápices. Maculae absent. Soralia
usually concolor to the thallus or more pale, linear continuous or interrupted, sometimes with a
tendeney to become extensive or labriform, or then orbicular in the apex of small lateral growths
(lacinules?), usually marginal, rarely laminai, in apothecial primordia; soredia granular.
Lacinules, isidia and pustules absent. Medulla white. Undersurface black, lustrous, smooth to
rugose or papillate, with scars; marginal zone brown, lustrous, 3-12 mm wide, naked, with an
attenuated limit, smooth, rugose or papillate, sometimes veined; rhizines black, simple, up to
1 mm long, frequent, distributed in groups.
Apothecia concave/ cupuliform, up to 3 mm in diameter, stipitate, laminai, margin sorediate,
amphithecium sorediate, disc dark brown, epruinose, imperforate; epithecium 10-12 um;
hymenium 30-34 um; subhymenium 12-14 um; ascospores ellipsoid, 25-28 x 15-18 um,
79
episporium 2-3 um. Pycnidia laminai, conspicuous, without prominent margin, few, ostiole
black; conidia not found (ca. 30 pycnidia investigated).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema sieberi is characterized by the sorediate thallus with wide lobes (8-25
mm), the simple to furcated cilia, up to 2 mm long.
It is very similar to P. cristiferum, the differences being the presence of well developed cilia,
wider lobes, soralia not so crisped and the absence of eumitrins. These features seem suitable for
species separation at the present time.
On the other hand, the ascospores are similar, and one of the specimens determined as P.
sieberi by Dodge (Mauritius, Wight s.n., FH!), actually the duplicate of the lectotype ofParmelia
cristifera, has sublageniform conidia, 5-7 x ca. 1 um, equal to that described for Parmelia
cristifera (see Vainio 1909a). The chemistry of the two species is the same.
If we consider P. sieberi as synonym of P. cristifera, we must do accept that Parmelia
cristifera could produce well-developed cilia. Krog & Swinscow (1981) already pointed out that
occasional, rudimentary cilia could grow in the lobe axils of P. cristifera. InP. sieberi, however,
the cilia are very well developed, seen by the unaided eye.
Krog & Swinscow (1981) described the holotype of P. sieberi as a collection from Wight, at
Mauritius (FH). Elix, in a label, did the same. The true holotype, however, is "Mauritius, Sieber,
Crypt Exot. 44, corticole, ex herb. Sbarbaro, at Farlow Herb.", as indicated in the protologue
(Dodge 1959), by a label from Dodge dated 1957 and another one from Kurokawa dated 1967.
There is also an isotype in FH, smaller than the holotype, but from the same place. It has very
few cilia and more crisped soralia, thus it is more adequately treated as Parmelia cristifera.
Unfortunately, pycnidia were not found.
The absence of pycnidia in the lectotype of Parmelia cristifera and in the holotype and
isotype of Parmelia sieberi leaves a gap, which can be filled, hopefully, with the study of fresh
collections.
The lectotype of Parmelia var. clemensiae Vainio (TUR!) agrees in gross morphology, and
conidia (sublageniform, 6-9 x ca. 1 um) with P. cristiferum, although presents also well
developed cilia and the chemistry is identical with that of P. sieberi, and so is accepted here as a
synonym of this last species.
Distribution: Africa (Dodge 1959) and Philippines (Vainio 1909b).
Parmotrema spinibarbe (Kurok.) Hale ex DePriest & B. Hale
Mycotaxon 67: 204. 1998.
Parmelia spinibarbis Kurok., Bulletin of the National Science Museum Tokyo 17 (4): 299. 1974.
- Type: Brazil, Rio de Janeiro, [Municipality of] Petrópolis, about 810 m alt, on trees, July 17,
1971, leg. S. Kurokawa 8348 (holotype: TNS!; isotypes: DUKE!, G!, O!, US!, W!).
(Figs. 69-71)
80
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, corticicolous, 8-1 1 cm broad. Lobes
irregularly branched, laterally overlapped, 3-15 mm wide, surface smooth to more often strongly
cracked, especially at old parts, rather opaque; apical zone ± rounded; margin crenate to usually
incised, undulated. Maculae usually strong, although difficult to see at the first sight, since it
covers almost ali the upper surface, punctiform to irregular, sometimes assuming a pseudo-
reticular pattern (by the coníluence of the dots), laminai, not originating cracks (as normally
occur with real reticular maculae). Cilia black, simple to more commonly cespitose, 0.5-1.5 x
0.02-0.1 mm, abundant, apparently friable, since the upper surface is abundantly covered by
pieces of cilia. Soralia marginal, somewhat extensive toward the centre of the thallus, or forming
both laminai and marginal botryose structures fully covered by soredia; soredia granular.
Lacinules, pustules and isidia absent. Medulla white. Undersurface black, lustrous, papillate or
rugose, with many cracks; marginal zone brown (sometimes absent), lustrous, 0.5-2.0 mm wide,
naked, with a sharp limit, smooth to slightly rugose or less frequently papillate; rhizines black,
simple, 0.3-1.5 x 0.02-0.10 mm, abundant, distributed in groups.
Apothecia plane or concave, 3-10 mm in diameter, shortly stipitate, laminai, margin
sorediate, amphithecium maculate, sorediate, disc dark brown, epruinose, perforate at maturity;
ascospores ellipsoid to broadly ellipsoid, 12.0-13.5 x 7.0-9.0 um, episporium 1-1.5 um.
Pycnidia submarginal, conspicuous, without or with a slightly prominent margin, abundant,
ostiole black; conidia filiform, 7.5-10.0 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV+ orange.
TLC in solvent C, HPLC (J.A. Elix, 17.VII.1996, label with the holotype in TNS): atranorin
(minor), chloro atranorin (minor), lichexanthone (major), salazinic acid (major), consalazinic acid
(trace).
Remarks: Parmotrema spinibarbe is characterized by the presence of cespitose cilia (Fig. 70),
and the marginal and less frequently laminai soredia, sometimes growing from botryose
structures (Fig. 71).
InP. pontagrossense , otherwise similar in the cilia branching, cortical chemistry and conidial
form and size, the botryose structures are absent from the examined type material (holotype,
UPCB, isotype in BAFC). Additionally, it produces lichesterinic and protolichesterinic acids in
the medulla. Unfortunately, apothecia are unknown in this species, and so we cannot compare
ascospores data.
Parmotrema diffractaicum (Essl.) Hale is morphologically very similar to P. spinibarbe, as
already noted by Fleig (1997) and Canêz (2005). Chemically, however, they are distinct since P.
diffractaicum produces diffractaic acid instead salazinic acid.
The holotype of P. spinibarbe (TNS) is in very good condition and has conidia and apothecia.
Ali other isotypes seen agree with the holotype in every aspect. However, some notes can be
added: (1) the botryose structures are much more abundant in the isotype from G; (2) the isotype
from O has the upper córtex damaged in some lobes that resulted in the production of many
orbicular atypical soralia; additionally, the marginal cilia are almost entirely cespitose; (3) a well
developed perforate apothecium and many botryose structures were observed in the isotype from
US that had already been examined by S. Eliasaro & M. Adler (label dated 1997); (4) the isotype
from W also developed abundant botryose structures and the upper córtex is injured in some
lobes, however the soralia is not so well developed.
Within the additional specimens examined, two of them do not developed the botryose
structures (Spielmann 1348 and Spielmann 1350), being probably young thalli. The latest
81
(Spielmann 1350) also has the ápices of some lobes pruinose. The specimen Spielmann 4920 is
33 cm long, the larger thallus found.
Distribution: known only for Brazil, being recorded to the States of Paraná, Rio de Janeiro
(Kurokawa 1974), Rio Grande do Sul (Fleig 1997, Canêz 2005, Spielmann 2005), Santa Catarina
(Fleig 1997) and São Paulo (Fleig 1997, Ribeiro 1998, Benatti 2005).
Additional specimens examined: Brazil, Rio Grande do Sul State, Municipality of Boqueirão
do Leão, Linha Sinimbuzinho, Perau da Nega, 29°20'17.2"S, 52°26'33.6"W, 430 m alt,
saxicolous, open place, near the waterfall., 23.11.2004, leg. A.A. Spielmann & L.S. Canêz 1348,
1349, 1350 (SP). The same, Municipality of Santa Cruz do Sul, near the Residencial Costa
Norte, 29°41'25.3"S, 52°26'19.3"W, 100 m ali, corticicolous, on forest border, 26.VI.2004, leg.
A.A. Spielmann 1168, 1171, 1172, 1200 (SP). The same, São Paulo State, Municipality of Mogi-
Mirim, Estação Experimental do Instituto Florestal, corticicolous, border of Cerradão vegetation,
14. V.2004, leg. A.A. Spielmann, M.P. Marcelli, P. Jungbluth & M.N. Benatti 4920.
Parmotrema stuppeum (Taylor) Hale
Phytologia28 (4): 339. 1974.
Parmelia stuppea Taylor, London Journal of Botany 6: 175. 1847. - Type: U.S. A, Califórnia,
Monterey, leg. Beechy [not "Beechey"] (holotype: FH!; isotypes: BM!, K).
(Fig. 72)
Holotype description
Thallus brownish (herbarium), lobate, apparently loosely adnate, 4 cm broad. Lobes irregularly
branched, crowded, 3-7 mm wide, surface extensively cracked; apical zone rounded; margin
smooth to dentate, undulated. Maculae apparently present in some áreas, barely distinct and
difficult to interpret because of the thallus state of preservation, maybe punctiform or effigurate.
Cilia absent? (Only something similar to one cilium was found, black, ca. 0.5 mm, simple).
Lacinules, pustules and isidia absent. Soralia marginal or extensive; soredia farinose.
Undersurface black and with abundant reddish brown stains, lustrous, strongly venate, naked,
with scars; marginal zone black or with brown stains, naked, with veined; rhizines black, seen
only through one hole in the thallus where somebody made a spot test (possibly Hale).
Apothecia plane or more usually concave, up to 7 mm in diameter, short-stipitate, laminai,
margin smooth, crenate or with deep incisions, sorediate, amphithecium usually sorediate, disc
dark brown, probably paler when freshly collected, epruinose, imperforate, often cracked;
ascospores not formed or immature. Pycnidia absent.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema stuppeum is recognized by the strictly marginal soralia, absent lacinules,
sublageniform conidia and the médium sized ascospores (12-17 x 6-9 um fide Hale 1965).
It is so very close to P. cristiferum, that has larger ascospores (24-35 x 12-20 um). Another
similar species isP. maximum, differentiated by the sorediate lacinules.
82
The holotype of P. stuppeum in FH is probably little representative of the species (see Hale
1965) and the same can be said about the isotype from BM. Ascospores and conidial data could
not be obtained. On the other hand, the information available about P. stuppeum is not
conclusive at the present time. Taylor (1847) do not provided microscopic data. The description
of Hale (1965) reported ascospores 12-17 x 6-9 um, episporium 1.0-1.5 um, and conidia 4-6
um. However, it is not clear what specimen Hale used to obtain these data, whether from the
type of P. stuppea or from the type of P. máxima (since that was the type illustrated) placed in its
synonymy, or from additional specimens examined.
There's a label from Berry (E.C.B), dated from July, 02, 1937, asserting that the specimen in
FH could be the type of Parme lia stuppea. However, in his publication (Berry 1941) he placed
Parmelia stuppea as belonging to Lecanora.
Some published pictures oíParmotrema stuppeum, like that of Hale (1979) and Brodo et ai.
(2001), show the strictly marginal soredia not occurring in lacinules. This concept agrees with
Hale (1965: 291), where he clearly stated that Parmelia stuppea lacks distinct lacinules (as
"laciniae").
Distribution: Africa (Hale 1965), Ásia (Divakar & Upreti 2005), Europe (Hale 1965), North
America (Hale 1965, Brodo et ai 2001) and Central America (Hale 1965).
Parmotrema subcaperatum (Kremp.) Hale
Phytologia28 (4): 339. 1974.
Parmelia subcaperata Kremp., Videnskabelige meddelelser fra Dansk naturhistoriske Forening i
Kjõbenhavn 5: 10. 1874. - Type: Brazil, [Minas Gerais State, Municipality of Caeté], Serra da
Piedade, ad truncos arborum, leg. Warming 297 (holotype: M!).
= Rimeliella subcaperata (Kremp.) Kurok., Armais of the Tsukuba Botanical Garden 10: 7.
1991.
= Canomaculina subcaperata (Kremp.) Elix, Mycotaxon 65: 477. 1997.
= Parmelia imperforata Nyl., Acta Societates Scientiarum Fennicae 26 (10): 7. 1899. Type:
Brazil, 1871, leg. Glaziou 1839 (holotype: H-NYL n° 35425).
(Fig. 73)
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, corticicolous, 4 cm broad. Lobes
irregularly branched, crowded, 3-7 mm wide, surface continuous, smooth to subfoveolate,
lustrous, becoming rugose, cracked and foveolate towards the center; apical zone rounded;
margin crenate to irregular or sublacinulate, plane to undulated. Maculae distinct or sometimes
weak, effigurate, laminai, originating cracks or not. Cilia black, simple and apparently also
rarely furcated, up to 1.5 mm long, quite thickened, 0.04-0.10 mm wide, few. Lacinules,
pustules, soredia and isidia absent. Medulla white. Undersurface black or brown blackish,
opaque, slightly rugose or papillate, with cracks; marginal zone dark brown or brown, opaque to
lustrous, 3-10 mm wide, naked or rhizinate, with an attenuated limit, smooth to papillate or
veined; rhizines black, dimorphic, the shorter ones simple, up to 1.0 mm long, thin, 0.02-0.05
mm wide, few, irregularly distributed, the longer ones up to 2 mm long, thickened, 0.05-0.10
mm wide, simple or irregularly branched near the ápices, frequent, irregularly distributed.
Apothecia cupuliform, up to 8 mm in diameter, substipitate, laminai or submarginal, margin
incised to lacerate, usually involute, amphithecium maculate, strongly rugose and foveolate
83
when mature, disc dark brown to blackish, epruinose, perforate at maturity; ascospores ellipsoid,
14-16 x 9-11 um, episporium 1.0-1.5 um. Pycnidia submarginal to less frequently laminai,
conspicuous, without prominent margin, abundant, ostiole black; conidia not found (ca. 30
pycnidia examined).
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
HPLC (J.A. Elix, 23. VIL 1997, label with the holotype): atranorin (minor), usnic acid (minor),
salazinic acid (major), consalazinic acid (trace).
Remarks: Parmotrema subcaperatum is characterized by the effigurate maculae, dimorphic
rhizines, absence of propagules and the widely black or brown blackish undersurface, with a
marginal and narrow dark brown área.
It is related to P. erubescens, already considered conspecific (Hale 1965, Winnem 1975,
Kurokawa 1991). However, P. subcaperatum has the undersurface black in a wide área, usually
slightly rugose to veined, the marginal zone being dark brown and the upper surface
subfoveolate to foveolate. On the other hand, in P. erubescens, the underside is blackish brown
only in the very center, smooth to papillate, the marginal área very light brown, and the upper
surface only scrobiculate.
The main characters accepted today for the separation of these species are presented in the
Table3.
Some problems still remain and could not be here clarified. The holotype of P. subcaperatum
is corticicolous and coriaceous, with ascospores 14-16 x 9-11 um and, although with abundant
pycnidia, conidia were not seen. The lectotype of P. erubescens is corticicolous,
membranaceous, without apothecia but with conidia 8-11 x ca. 1.0 um. The features of the
underside seems to be comparable to the characters listed by Elix (1997) and so we are accepting
these species as good taxa, at least up to more material become available for study.
Table 3. Main characters of P. subcaperatum andP. erubescens (compiled from Elix 1997)
P. subcaperatum P. erubescens
Substrate saxicolous or rarely corticicolous corticicolous
Thallus consistence membranaceous coriaceous
Habitat temperate subtropical and tropical
Lower surface black to the margins or marginal with a broad and brown marginal
zone dark brown, narrow and naked zone, papillate or with short, fine
rhizines
Fine rhizines 0.5-1.0 mm long 0.05-0.1 mm long
Coarse rhizines solitary, rare grouped in the center
Ascospores 12-19 x 6-10 um 10-12 x 8-10 um
Conidia 9-14 x 1 um 10-14 x l ^ m
It must be noted that the holotype of P. subcaperatum, although full of apothecia, is devoid of
several entire lobes, and some features, like lobe margins, cilia, etc, could not be well evaluated.
The rhizines also seem not so clearly dichotomous, probably because the absence of the lobe
ápices, where they are normally more common.
The holotype oíParmelia imperforata is well developed, with ascospores 11-16 x 10-12 um,
episporium ca. 1 um and conidia 8-10 x ca. 1 um. The underside is paler that P. subcaperata,
84
and we are accepting it here as synonym with certain reserve, since this feature could not be
adequately studied in the holotype of P. sub cape ratam. Chemically, the presence of usnic acid
was not detected, the holotype having atranorin (minor), chloroatranorin (trace), salazinic acid
(major) and consalazinic acid (minor).
Distribution: Oceania and South America (Kurokawa 1991). In South America it is known to
Argentina (Calvelo & Liberatore 2002), Brazil (Zahlbruckner 1930a, Kurokawa 1991, Marcelli
2008), Paraguay (Osório 1970), Uruguay (Osório 1972) and Venezuela (Vareschi 1973). In
Brazil it was recorded to the States of Minas Gerais (Kurokawa 1991), Mato Grosso do Sul
(Osório 1992a), Paraná (Osório 1977b, Kurokawa 1991), Rio de Janeiro (Kurokawa 1991), Rio
Grande do Sul (Spielmann 2005, 2006) and São Paulo (Kurokawa 1991, Marcelli & Benatti
2008).
Parmotrema subis idiosum (Múll. Arg.) Hale
Phytologia 28 (4): 339. 1974.
Parmelia cetrata var. subisidiosa Miill. Arg., Engler Botanische Jahrbucher 20: 256. 1894. -
Type: Africa, Bumba, Usambara, leg. Holst 8772 p.p. (lectotype: G!), fide Hale & Fletcher
(1990).
= Parmelia subisidiosa (Miill. Arg.) Dodge, Armais of the Missouri Botanical Garden 46: 87.
1959.
= Rimelia subisidiosa (Miill. Arg.) Hale & Fletcher, The Bryologist 93 (1): 29. 1990.
(Fig. 74)
Holotype description
Thallus brownish, lobate, apparently loosely adnate, corticicolous with mosses, 5.5 cm broad.
Lobes irregularly branched, laterally overlapped , 3-7 mm wide, surface continuous or cracked,
smooth or irregularly subscrobiculate, lustrous, becoming strongly cracked and irregular towards
the center; apical zone rounded; margin smooth to crenate, undulated. Maculae distinct, reticular,
laminai. Cilia simple, short, up to 0.6 mm long, 0.02-0.05 mm wide, few. Lacinules, pustules
and soredia absent. Isidia concolor with the thallus, apex usually darker, simple to coralloid,
0.1-0.2 x 0.05-0.15 mm, erect, firm, or less frequently flaking off, sometimes ciliate, laminai or
marginal. Medulla white. Undersurface black, lustrous, smooth or slightly rugose, with few
cracks; marginal zone dark brown, pale brown or variegating these colors, lustrous, 1-2 mm
wide, naked, with an attenuated limit, smooth or papillate; rhizines black, simple, irregularly
branched or squarrose, sometimes with flatted ápices, up to 2 mm long, abundant, evenly
distributed.
Apothecia absent. Pycnidia submarginal, inconspicuous, without prominent margin, rare,
ostiole black; conidia not found.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
85
Remarks: Parmotrema subisidiosum is characterized by the formation of simple to coralloid
isidia, the reticular maculae and the presence of salazinic acid.
Parmotrema lacteum Marcelli & Spielmann, a close species recently described (Marcelli et
ai. 2007), is distinguished by the formation capitate sorediose clusters originating isidioid
soredia, and the milky thallus (see comments under this species).
Another species with reticular maculae and isidia is P. bonplandii, distinct by the chemistry:
norlobaridone and lichexanthone instead salazinic acid (Mata Garcia 1994).
The holotype of P. subisidiosum is well-preserved, while glued by the underside to a paper.
The observations of this part were made in a free part. Unfortunately, it doesn't have apothecia,
and conidia were not found in the rare pycnidia seen.
There's a label from H. des Abbayes (dated from 1958), giving the spot tests and providing a
new combination, "Parmelia subisidiosa (Miill. Arg.) des Abb. comb. nov.". A second label
made by Hale in 1959, gives the combination as "P. subisidiosa (Miill. Arg.) Dodge". In fact,
Dodge (1959) published the new combination, and apparently, Abbayes did not.
Distribution: Africa (Dodge 1959, Krog & Swinscow 1981, Hale & Fletcher 1990), North
America (Moore 1968, Dey 1978, Brodo et ai 2001), Central America (Tenório et ai. 2002) and
Tropical America (Hale & Fletcher 1990), where it was recorded to Brazil (Marcelli 2008),
Uruguay (Osório 1992b) and Venezuela (Feuerer 2008). In Brazil it was reported to the States of
Rio Grande do Sul, Santa Catarina (Fleig 1997), São Paulo (Marcelli 1991, Ribeiro 1998, Benatti
2005).
Parmotrema subsumptum (Nyl.) Hale
Mycotaxon 5 (2): 434. 1977. *
Parmelia subsumpta Nyl., Flora 52: 1 17. 1869. - Type: Brazil, Minas Gerais State, corticicolous,
leg. Glaziou s.n. (holotype: H-NYL n° 3545 1 ! )
= Rimeliella subsumpta (Nyl.) Kurok., Armais of the Tsukuba Botanical Garden 10: 9. 1991 .
= Canomaculina subsumpta (Nyl.) Elix, Mycotaxon 65: 477. 1997.
= Parmelia urceolata Eschw. var. nuda Miill. Arg., Flora 63: 266. 1880. Type: Brazil, Rio de
Janeiro, prope [Municipality of] Petrópolis, leg. Deventer 33 (holotype: G).
= Parmelia hypotropa var. imperialis Hue, Nouvelles Archives du Muséum d'Histoire Naturelle
de Paris, ser. 4 (1): 189. 1899. Type: Brazil, corticicolous, 1887, leg. S.M. Dona Theresa
Christina Maria, imperatrix (holotype: P).
= Parmelia corrugis (Fr.) Miill. Arg. var. imperialis (Hue) Zahlbr., Catalogus lichenum
universalis 6: 236. 1930.
= Parmelia leucoxantha Miill. Arg. f firma Sambo, Annali di Botânica 22 (1): 18. 1939. Type:
Brazil, Rio de Janeiro, Serra dos Órgãos, May 1839, leg. Casaretto 2444 (holotype: RO), fide
Hale (1965).
(Fig. 75)
Holotype description
Thallus brownish (herbarium), lobate, ± adnate, corticicolous, 8 cm broad. Lobes irregularly
branched, crowded, usually strongly crisped, 2-10 mm wide, surface usually irregularly cracked,
smooth or with irregular depressions, lustrous, becoming reticulately cracked towards the center;
86
apical zone rounded; margin sinuous-incised, usually strongly crisped, sometimes undulated.
Maculae weak, effigurate, laminai, originating cracks. Cilia black, simple, up to 1.5 mm long,
0.02-0.05 mm wide, rare. Lacinules , pustules, and isidia absent. Soralia white, growing from the
margin of the lobes and becoming extensive towards the undersurface, the sorediate lobe
margins becoming revolute and strongly crisped; soredia farinose to subgranular. Medulla white.
Undersurface dark brown to brown, blackening only in the very centre, sublustrous, smooth to
papillate or slightly veined, with sparse cracks; marginal zone absent (brown) or pale brown,
then 1-5 mm wide, lustrous, naked or rhizinate, with a sharp limit, smooth to veined or with very
slight depressions; rhizines black, dimorphic, the shorter ones simple or sometimes squarrose,
usually curled, up to 0.5 mm long, thin, 0.01-0.04 mm wide, abundant, evenly distributed; the
longer ones up to 3.5 mm long, thickened, 0.05-0.12 mm wide, simple or more commonly
irregularly branched, especially near the apex, frequent, distributed in groups.
Apothecia cupuliform, 1.3-3.2 mm in diameter, stipitate, laminai, margin smooth to incised,
usually involute, sorediate or not, amphithecium maculate, smooth to irregular, disc pale brown
to dark brown, epruinose, perforate; ascospores not seen (11-14 x 7-9 um fide Nylander 1869).
Pycnidia submarginal, conspicuous, without prominent margin, few, ostiole black; conidia not
found (ca. 20 pycnidia examined) see comments below under P arme lia arceolata var. nuda.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema subsumptum is characterized by the strongly crisped lobes sorediate at
the undersurface, the effigurate (although weak) maculae, the dichotomous rhizines and the
simple and rare cilia. The ontogeny of the soralia is quite interesting, since they start at the
margin and extends backwards on the undersurface, not the upper surface as is common in
Parmotrema.
In P. petropoliense, before thought to be a synonym (Hale 1965), the soralia are formed in the
margin or on the upper surface of conspicuous lacinules that form pustuloid structures (i.e., not
extensive). Additionally it has abundant, long (up to 3 mm) and usually furcated cilia.
Parmotrema leucosemothetum can be distinguished by the soralia almost exclusively
marginal, the strongly maculate upper córtex (with effigurate maculae), the simple, usually
thickened and ± tapered cilia, and the distinctly shiny bare marginal zone on the undersurface.
Finally, P. margaritatum has the soralia growing subapically in lacinules, the upper surface
hypermaculate, and a black undersurface with monomorphic rhizines.
The holotype of P. subsumpta seems to have been pressed, as can be seen by the strongly
ílattened lobes. It has three poor developed apothecia, and one tentative search by ascospores
was made in the larger one. However, the hymenium was immature and no ascospores were
found. So we have to trust in the measurements given by Nylander. However, he published
(Nylander 1869) ascospores with 11-14 x 7-9 um, although his handwriting on the envelope
states "14-16 x 7-9 um". Hale (1965) found ascospores 12-15 x 7-10 um. Analyses of more
specimens can help to establish the real range of ascospores variation in this species.
The holotype oíParmelia urceolata var. nuda agrees well withP. subsumpta. It is completely
eciliate, and the ontogeny of the soralia described for P. subsumpta is more evident here. Conidia
were found and they are filiform, 7.5-12 x ca. 1.0 um. The information concerning the collector
(Deventer 33) could not be confirmed since there is no label about this. So this datum comes
from Hale (1965).
Parmelia hypotropa var. imperialis Hue has as holotype three thalli, ali of them agreeing with
Parmotrema subsumptum in the overall morphology, being scarcely ciliate. One thallus has
several well developed apothecia, up to 6 mm in diameter, laminai, substipitate to stipitate,
87
usually strongly maculate and longitudinally rugose, disc perforate, the margin irregularly
sorediate, the ascospores (most immature) being 14.0-16.5 x 7-9 um. It has the undersurface
extensively dark brown and black in some áreas while in the larger thallus (13 cm broad, lobes
5-18 mm wide) it is entirely brown. Few pycnidia are present is this thallus, but no conidia was
found (ca. 20 pycnidia examined). The third thallus seems to be part of the fertile thallus. Hue's
(1899) description is a combination of the features from the three thalli. The two smaller thalli
are undoubtedly P. siibsumptum. The larger thallus is maybe a different species.
Finally, the holotype ofParmelia leucoxantha f. firma is also identical with the holotype of P.
subsumpta, only the undersurface was not studied in detail since the thallus is strongly attached
to a bark.
Distribution: Africa (Hale 1965, Winnem 1975, Krog & Swinscow 1981), Ásia (Kurokawa
1991, Kurokawa & Lai 2001, Chen et ai 2003, Divakar & Upreti 2005) Oceania (Kurokawa
1991, Elix 1994, Louwhoff & Elix 1999) and Américas (Hale 1965, Kurokawa 1991). In South
America it is known for Argentina (Hale 1965, Kurokawa 1991), Brazil (Hue 1899), Uruguay
(Osório 1979, Kurokawa 1991) and Venezuela (Hale 1965). In Brazil was reported to the States
of Minas Gerais (Nylander 1869, Hale 1965, Kurokawa 1991, Ribeiro 1998), Paraná (Eliasaro
2001), Rio de Janeiro (Míiller Argoviensis 1880, Cengia Sambo 1939c, Kurokawa 1991), Rio
Grande do Sul (Spielmann 2006) and São Paulo (Hale 1965, Ribeiro 1998).
Parmotrema subtinctorium (Zahlbr.) Hale
Phytologia28 (4): 339. 1974.
Parmelia subtinctoria Zahlbr., in Handel-Mazetti, H.: Symbolae sinicae. Botanische Ergebnisse
der Expedition der Akademie der Wissenschaften in Wien nach Sudwest-China 1914/1918, III,
Lichenes. J. Springer, Wien, p. 193. - Type: China, Prov. Yílnnan, prope vicum Sanyingpan ad
septentr. urbis Ytinnanfu, 26° lat, in regione calide temperata, substr. córtice Quercuum, alt. s.m.
ca. 2.400 m, 14.111.1914, leg. Dr. Heinr. Frh. v. Handel-Mazzetti 5645 (Diar. Nr. 146) (holotype:
WU!).
= Rimeliella subtinctoria (Zahlbr.) Kurok., Armais of the Tsukuba Botanical Garden 10: 10.
1991.
= Canomacidina subtinctoria (Zahlbr.) Elix, Mycotaxon 65: 477. 1997.
(Fig. 76)
Holotype description
Thallus brownish (herbarium), lobate, loosely adnate, corticicolous, 10 cm broad. Lobes
irregularly branched, laterally overlapped to crowded, usually strongly curled, 4-12 mm wide,
surface slightly scrobiculate, becoming strongly cracked in the center; apical zone and margins
usually curled down, when visible the apical zone is rounded, and the margin crenate to incised,
plane. Maculae distinct, effigurate, laminai. Cilia black, simple, up to 1 mm long, 0.02-0.05 mm
wide, few and usually broke. Lacinules, pustules and soredia absent. Isidia concolor with the
thallus or little paler, apex dark brown, simple to coralloid, 0.1-1.0 x 0.05-0.2 mm, erect, firm,
apex eciliate, laminai. Medulla white. Undersurface dark brown to black in the central zone,
lustrous, slightly rugose or papillate, with scars; marginal zone brown to pale brown, lustrous, 2-
5 mm wide, naked or with initial rhizines, with an attenuated limit, difficult to distinguish,
slightly rugose and papillate, except in some áreas where remains smooth; rhizines black or dark
brown, abundant, dimorphic, the smaller ones simple and usually curled, up to 0.5 x 0.05 mm,
more or less evenly distributed, the larger ones simple to sometimes branched, up to 3.0 x 0.2
mm, distributed in groups.
Apothecia absent (according to Hale 1965, they are rare, with amphithecium isidiate and the
disc imperforate, ascospores 8-11 x 5-8 um). Pycnidia submarginal, conspicuous, without
prominent margin, few, ostiole black; conidia not found (ca. 30 pycnidia examined.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), norlobaridone (minor), loxodin (trace), protocetraric acid (trace).
Remarks: Parmotrema subtinctorium is distinguished by the isidiate thallus with effigurate
maculae, the undersurface slightly rugose to papillate, the absence of usnic acid and the presence
of norlobaridone.
It is morphologically cio se to P. neotropicum, which isidia can become sorediate, the
underside is abundantly veined and additionally has usnic acid and lacks norlobaridone.
Kurokawa (1991) reduced Parmotrema haitiense (Hale) Hale to the synonymy of P.
subtinctorium. This species was not studied here, since it lacks salazinic acid, but we agree with
Elix (1993) and Fleig (1997) that they are distinct species.
Hale (1965) reported some synonyms to P. subtinctorium, but they were still not investigated:
Parmelia virens Mull. Arg. f. isidiosa Mull. Arg., Parmelia leucosemotheta Hue f. isidiata Hue,
Parmelia velutina Zahlbr. [non Parmelia velutina (A eh.) Wallr.], and Parmelia protovirens
Gyelnik.
Distribution: Africa (Hale 1965, Winnem 1975, Krog & Swinscow 1981), Ásia (Hale 1965,
Park 1990, Kurokawa 1991, Kurokawa & Lai 2001, Chen et ai 2003, Divakar & Upreti 2005)
and Américas (Hale 1965). In South America it is known to Paraguay (Hale 1965). In Brazil it
was recorded to the States of Minas Gerais (Ribeiro 1998) and Paraná (Donha 2005).
Parmotrema ulíralucens (Krog) Hale
Mycotaxon 1 (2): 108. 1974.
Parmelia ultralucens Krog, The Bryologist 77 (2): 253. 1974. - Type: U.S. A, Geórgia, Harris
Co., DowdelTs Knob, on rock, 1965, leg. H. Krog (holotype: O!; isotypes: BM!, S and US).
= Canomaculina ultralucens (Krog) Elix & J.B. Chen, Mycotaxon 86: 21. 2003.
= Parmelia suberinita auet, non Nyl.
= Parmelia monumentalis Asahina (not validly published, fide Krog 1974, p. 256).
(Fig. 77)
Holotype description
Thallus pale brownish (herbarium), lobate, loosely adnate, saxicolous, 18 cm broad. Lobes
irregularly branched, laterally overlapped, 3-1 1 mm wide, surface continuous, smooth to
irregular, lustrous, sometimes pruinose in the ápices, becoming rugose toward the center; apical
zone rounded; margin crenate, undulated. Maculae weak and inconspicuous, although covering
mo st of the surface, punctiform to effigurate, usually forming a pseudo -reticular pattern. Cilia
black, simple or cespitose, 0.5-1.0 (-1.3) x 0.02-0.05 mm, abundant. Isidia paler or darker than
89
the córtex, apex strongly dark brown, simple to less frequently branched, sometimes coralloid,
up to 0.5 x 0.05-0.10 mm, erect, firm, apex sometimes ciliate laterally, laminai and marginal.
Lacinules, soralia, and pustules absent. Medulla white. Undersurface black, lustrous, smooth to
papillate or slightly rugose, with cracks; marginal zone dark brown, lustrous, 1.5-4.0 mm wide,
naked, with an attenuated limit, smooth to slightly rugose; rhizines black, simple or rarely
branched, up to 1.5 mm x 0.01-0.10 mm, abundant, evenly distributed.
Apothecia absent (according to Krog 1974, apothecia rare, up to 8 mm in diameter, disc
imperforate, ascospores 15-17 x 10-12 um). Pycnidia submarginal, conspicuous, with or
without prominent margin, few, ostiole black; conidia short filiform, 6-10 x ca. 1 .0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > orange red, C-, KC-, P+ strong
yellow, UV+ yellow.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor), lichexanthone (minor).
Remarks: Parmotrema ultralucens is characterized by the isidiate thallus, the cespitose cilia and
the presence of both lichexanthone and salazinic acid in the medulla.
When describing the species, Krog (1974) asserted that it has rare apothecia. However, no
apothecia at ali were found in the holotype, and she does not indicate from what specimen the
apothecia examined belong. The isotype from BM agrees well with the holotype but it also lacks
developed apothecia.
Krog (1974) also recorded that Hale (1965) selected as neotype to Parmelia subcrinita Nyl.
the specimen H-NYL 35472; however, the specimen listed by Hale actually is H-NYL 35479. If
they are different species or this was a typographical error remains an open question.
The name Parmelia subcrinita, although rejected by Krog (1974) according to the Article 69
of the Seattle Code (Stafleu et ai 1972), was still not formally proposed for rejection.
Distribution: Africa (Krog 1974, Swinscow & Krog 1988), Ásia (Hale 1965, Krog 1974,
Kurokawa & Lai 2001, Elix et ai 2002, Chen et ai 2003), Azores (Hale 1965, Krog 1974),
Oceania (Hale 1965, Krog 1974, Elix 1994, Louwhoff & Elix 1999), North America (Hale 1965,
Esslinger 2008), Central America (Hale 1965), and South America (Hale 1965, Krog 1974). In
South America it was recorded to Brazil, Colômbia and Ecuador (Hale 1965). In Brazil it was
cited for the States of Minas Gerais (Hale 1965), Rio Grande do Sul (Spielmann 2006), Santa
Catarina (Marcelli 1992) and São Paulo (Marcelli 1992, Benatti 2005).
Parmotrema verrucisetosum Sipman
Mycotaxon44(l): 8. 1992.
Type: Guyana, Upper Mazaruni district, trail from Kamarang river to Pwipwi mountain, N of
Waramadan, rocky savannah, c. 10 Km N of Waramadan, 05°57'N, 60°45'W, alt. ca. 800 m, 28
February 1985, leg. H. Sipman & A. Aptroot 19478 (holotype: B!).
(Figs. 78-79)
Holotype description
Thallus greenish-gray to whitish-straw towards the centre, lobate, loosely adnate, corticicolous,
3.0-7.3 (-10) cm broad. Lobes irregularly branched, laterally overlapped, 2-11 mm wide,
surface continuous, smooth, reticulately cracked towards the centre, opaque or slightly lustrous
in some áreas; apical zone rounded; margin crenate, plane or undulate. Lacinules absent.
90
Maculae absent. Cilia black, simple, furcated, dichotomous or sometimes cespitose, 0.5-3.5 x
ca. 0.05 mm, abundant, present in the margin and on the laminai and marginal warts. Pastiães,
soredia and isidia absent. Warts conical or ílatted, 0-2 x 1-4 mm, laminai and marginal, often
with one or more cilia. Medulla white, slightly orange in some áreas, probably due to the
decomposition of the salazinic acid. Undersurface black, lustrous, often smooth and with many
cracks, sometimes rugose or papillate, rarely scrobiculate; margin absent, beige or eburneous,
lustrous, 1-4 mm wide, naked, limit to attenuate to sharp, smooth, sometimes slightly rugose or
with barely distinct depressions; rhizines black, cylindrical or rarely flattened in the basal
portion, simple, 0.5-3.0 x 0.05-0.2 mm, frequent, distributed in groups.
Apothecia and pycnidia unknown.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solventA and B: atranorin and salazinic acid (Sipman & van Aubel 1992).
Remarks: Parmotrema vemicisetosum is characterized by the presence of (often) ciliate warts,
which distinguish it from ali other known Parmotrema.
The cilia, on sight, seem to have numerous lateral branches (bristles). With more attention,
however, one can perceive the bristles are just fungi growing on the cilia, sometimes reaching
the thallus surface. Actually, these are anamorphic fungi, Hyphomycetes (R.A.P Grandi, pers.
comm.). The under córtex presents several cracks, a feature already observed in several other
parmotremoid lichens with salazinic acid. Sipman & van Aubel (1992) described this in the
protologue.
When comparing the holotype with the protologue, the first item, thallus size, is a little
problematic. The holotype is composed by several pieces, one of them well developed. There is
no indication if ali were part of the same thallus, and even connecting the parts, the total size do
not reach 10 cm. In the protologue the size is reported as "up to at least 12 cm". This left us to
compare the holotype with the picture in the protologue, in which the lichens seems to be entire.
And it really is. However, it is not the picture of the holotype {Sipman & Aptroot 19478), but an
additional specimen studied {Sipman 26595, B, VEN), from Venezuela, not Guyana, collected in
1990. The warts detail is also from this specimen.
Distribution: Guyana, Venezuela (Sipman & van Aubel 1992).
Parmotrema warmingii (Vainio) Spielmann & Marcelli, comb. nov.
Parmelia warmingii Vainio, Acta Societatis pro Fauna et Flora Fennica 7 (1): 41. 1890. - Type:
based on P. angustata Kremp. non Pers.
= Parmelia angustata Kremp., Videnskabelige meddelelser fra den Naturhistoriske forening i
Kjõbenhavn 5: 13, 1873. Not P. angustata Pers. in Gaudichaud-Beaupré, Voyage autour du
monde, execute sur les corvettes 1'Uranie et la Physcienne, sur les ordres du Cap. L. de
Freycinet, en 1817-1820, Botanique, p. 195, 1826 [= Pannoparmelia angustata (Pers.)
Zahlbr., fide Galloway (2007)]. - Type: Brazil, Minas Gerais State, [Municipality of Caeté],
Serra da Piedade, leg. Dr. Warming 324 (lectotype: M!; duplicates of the lectotype: C!, W!).
(Fig. 80)
91
Lectotype description
Thallus dirty beige to blackened (herbarium), laciniate, loosely adnate, saxicolous, 4 cm broad.
Laciniae dichotomous, crowded, 0.5-2.0 mm wide, surface continuous to irregularly scared,
smooth, lustrous, becoming irregularly or reticulately cracked towards the center; apical zone
usually acute, sometimes truncate; margin smooth, with a conspicuous black line, plane to
slightly undulated. Maculae extensive (hypermaculate), laminai, not originating cracks. Cilia
black, simple, up to 2.3 mm long, 0.01-0.05 mm wide, abundant. Lacinules, soredia and isidia
absent. Medulla white. Undersurface black, lustrous, papillate to rugulose, with cracks; marginal
zone dark brown, lustrous, 0.5-2.5 mm wide, rhizinate, with an attenuated limit, smooth; rhizines
black, simple, usually entangled, up to 1.5 mm long, usually thin, rarely thickened, 0.01-0.15
mm wide, frequent, distributed in groups.
Apothecia cupuliform, 1.0-2.5 mm in diameter, sessile to substipitate, laminai or
submarginal, margin incised, amphithecium maculate, smooth to irregular, disc dark brown,
epruinose, imperforate; ascospores ellipsoid, 15-16 x 6-8 um, episporium 1.0-1.5 um. Pycnidia
laminai, conspicuous, without prominent margin, abundant, ostiole black; conidia short-filiform,
6.5-10 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
Chemistry: atranorin (minor), chloroatranorin (trace), salazinic acid (major), consalazinic acid
(minor).
Remarks: Parmotrema warmingii is characterized by the laciniate thallus with long cilia (up to
2.3 mm) and the hypermaculate upper surface.
It resembles P. herrei, a species here recognized and that can be differentiated by the
branched, thickened cilia and the reticulate maculate upper surface (not hypermaculate).
Another interesting feature of this lichen is the laminai distribution of the pycnidia. They are
almost universally submarginal in the other species of Parmotrema s.l. with salazinic acid.
The name Parmelia angustata (Krempelhuber 1873) is illegitimate, since is antedated by
Parmelia angustata Persoon (in Gaudichaud-Beauprél826) that is now Pannoparmelia
angustata (Pers.) Zahlbr. (Galloway 2007). Vainio (1890), aware of this situation, proposed the
new name Parmelia warmingii Vainio to the taxon he recognized as good. This name was then
used by Zahlbruckner (1902, 1909) to Brazilian lichens he studied, and was also listed in the
Catalogus lichenum universalis (Zahlbruckner 1930a).
Although Krempelhuber, Vainio and Zahlbruckner recognized this species, it was included by
Hale & Fletcher (1990) as a synonym o f Rime lia cetrata, based on a too wide species concept
that included several taxa. Parmotrema cetratum (lectotype: H-NYL!) is represented by a
laciniate thallus with distinct reticular maculae (not hypermaculate) and with shorter cilia, up to
1.3 mm long.
The lectotype of P. warmingii (M) is well-preserved and agreeing totally with the duplicates
of the lectotype from C and W, ali being probably part of the same collection. The W duplicate
has the more developed apothecia, one of them perforate. One specimen studied by Vainio
(Lichenes brasiliensis exsiccati 1164, M) also has an imperforate apothecium. Therefore, the
perforation seems to occur only in well-developed old apothecia.
Recent collections represented by more developed thalli (up to 10 cm broad) support the
differential feature s of P. warmingii, mainly the laciniate thallus with long cilia, the
hypermaculate upper surface and the well developed black marginal line in the laciniae. Some
specimens has little wider laciniae, like Canêz 2464 (up to 3.5 mm wide). The specimen
Spielmann 2044 has well developed perforate apothecia.
92
This species is known to Minas Gerais and São Paulo States in Brazil, restricted to highlands
(above 1200 m). However it is possible that it occurs in a wider área, since it could be named as
P. cetratum up to now.
Distribution: Brazil, Minas Gerais State (Krempelhuber 1873, Vainio 1890, Zahlbruckner 1902,
Hale & Fletcher 1990).
Additional specimens examined: Brazil, Minas Gerais State, [Municipality of Catas Altas],
Caraça, 1885, Vainio Lichenes brasiliensis exsiccati 1164 (M). Idem, at "Cascudos",
20°05'38.7"S, 43°30'00.3"W, 1300 m, saxicolous, at the margin of the stream, partial sun,
27.111.2006, AA. Spielmann, L.S. Canêz & M.P. Marcelli 2009, 2017 (SP). Idem, at "Trilha para
Capelinha", 20°05'47.6"S,43°29'06.4"W, 1340m, saxicolous, shaded by a tree, AA Spielmann,
L.S. Canêz & M.P. Marcelli 2044 (SP), L.S. Canêz, AA. Spielmann & M.P. Marcelli 1550 (SP),
.Idem, at "Tanque Grande", 20°06'10.1"S, 43°29'38.4"W, 1275 m, saxicolous, slightly shaded,
04.IV.2006, AA. Spielmann, L.S. Canêz & M.P. Marcelli 2171 (SP). Brazil, São Paulo State,
Municipality of Piquete, Pico dos Marins, next to the first massif, 22°30'30.8"S, 45°07'46.4"W,
1.900 m, saxicolous, open place, 27.1.2007, L.S. Canêz, D.F. Peralta, J. Bordin & J. Atwood
2464 (SP).
93
Nomina inquirenda
Parmelia badia Pers.
in Gaudichaud-Beaupré, Voyage autour du monde, execute sur les corvettes 1'Uranie et la
Physcienne, sur les ordres du Cap. L. de Freycinet, en 1817-1820, Botanique, p. 198. 1826. -
Type: Hawaiian Islands, Gandichand 78 (lectotype: PC!; duplicates of the lectotype: BM, G).
(Fig. 81)
However well preserved and not small, the lectotype (PC) is represented by a possibly burnt ill
prepared thallus (like Parmelia maura, se below), without apothecia or pycnidia. It has very
similar sorediate lobes to P. cristiferum, and the chemistry needs to be revisited. If it proves to
have salazinic acid, the nameP. badia takes precedence over Parmelia cristifera.
In the sheet of the lectotype of P. badium (P) there is a label indicating the number 79 and
'Parmelia flexuosa Delise, P. badia Pers.". The name P. flexuosa was not found in any
catalogue (Zahlbrucknerl934, Lamb 1963, Index Fungorum 2009) and was possibly never
published.
Parmelia cetrata Ach. f. sorediifera Vainio
Acta Societatis pro Fauna et Flora Fennica 7 (1): 40. 1890. - Type: Brazil, Minas Gerais State,
Sitio [now António Carlos Municipality], leg. Vainio, Lichenes Brasilienses Exsiccati 616
(lectotype: TUR!).
This taxon is represented by a small (2.5 cm broad) not well developed thallus without apothecia
and pycnidia growing with a fertile Hypotrachyna (Vainio) Hale. It has short open lacinules,
sorediate at the ápices. The soralia also develop submarginally, turning the lobes concave. This is
the pattern we found in many Brazilian specimens we named previously Parmotrema
clavuliferum (Spielmann 2005, Benatti 2005, Jungbluth 2006) and, in respect with the
submarginal soralia and short, open lacinules, they are clearly different from the type of
Parmelia clavulifera. Conidial and ascospores features could not be studied to clarify the status
of this taxon.
Parmelia cineras cens var. saxicola Rãs.
Suom. Eláin.Kasv. Seuran Vanamo 2: 45. 1947. - Type: Uruguay, Pan de Azucar, Maldonado,
Lamb (lectotype: H), fide Hale & Fletcher (1990).
The type was not sent by H. This taxon was listed as synonym of P. cetratum by Hale & Fletcher
(1990).
94
Parmelia cristifera Taylor f. cinerata Zahlbr.
Repertorium species novarum Fedde 33: 58. 1934. - Type: Formosa, Kuwarus, leg. Asahina 51
(holotype: W; isotype: BPI), fide Hale (1965).
The holotype was not sent by W and BPI could not be contacted.
Parmelia gossweileri C.W. Dodge
Armais of the Missouri Botanical Garden 46: 153. 1959. - Type: Angola, Chiloango, le£
Gossweiler 809 le (holotype: K), fide Hale (1965).
The holotype could not be found in K.
Parmelia imerinensis C.W. Dodge
Annals of the Missouri Botanical Garden 46: 143. 1959. - Type: Madagáscar, East Imerina, leg.
Hildebrandt (holotype: FH!).
Holotype description
Thalhis beige (herbarium), probably greenish-gray when fresh, lobate, loosely adnate,
corticicolous, 16 cm broad. Lobes irregularly branched, laterally overlapped, 5-13 mm wide.,
surface usually smooth, lustrous, extensively damaged and finely rugose; apical zone rounded;
margin smooth or crenate, undulated. Cilia absent (three small formations seen, but probably
rhizines). Lacinules, maculae, isidia and pustules absent. Soralia concolor to the thallus or
whitish, usually linear and marginal, less frequently capitate and growing in the lobe crests;
soredia granular. Medulla white. Undersurface black, lustrous, smooth or slightly rugose, with
scars; marginal zone brown, lustrous, up to 6 mm wide, naked, with an attenuated limit, smooth,
rugose or papillate; rhizines black, simple (few observed).
Apothecia and pycnidia absent.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C, HPLC (J.A. Elix, 03.XI.1998): atranorin (minor), chloroatranorin (trace),
salazinic acid (major), consalazinic acid (minor), methyl pseudo salazinate, unknowns
(minor/trace).
Remarks: Parmelia imerinensis was proposed by Dodge (1959) based on a peculiar cortical
structure. The holotype is represented by a thallus with the upper córtex extensively damaged,
eroded, that give it an appearance of finely rugo se/granulo se. In one área, the surface seems
maculated, but has no maculae in the rest.
At present time is difficult to compare anatomical information relating Parmeliaceae, since
very few investigations are available. Recent discoveries (Barbosa 2009) show that cortical
anatomy could be quite a powerful tool to taxonomy.
Since apothecia and pycnidia were not found, the status of P. imerinensis remains unclear.
There is a chemical difference between P. imerinensis and P. cristifera, since the last contains
eumitrins. Morphologically, there are much more laminai soralia in P. imerinensis, and the
marginal ones are not as crisped as in P. cristifera. On the other hand, cilia are absent, making
this species not conspecific withP. sieberi.
95
The holotype is glued by the underside, so the observations were limited to the free margins.
One of the specimens determined by Dodge (label dated 1957), is clearly another species,
since it do not has salazinic acid (medulla K+ slowly yellowish), but probably protocetraric acid
(Hildur Krog's label, TLC, 19..). Actually, this is the holotype of P arme lia paxinoides C.W.
Dodge (see Dodge 1959). The data from the protologue agree very well with the specimen.
Parmelia maura Pers.
in Gaudichaud-Beaupré, Voyage autour du monde, execute sur les corvettes 1'Uranie et la
Physcienne, sur les ordres du Cap. L. de Freycinet, en 1817-1820, Botanique, p. 198. 1826. -
Type: Hawaiian Islands, Gaiidichand (lectotype: L!; duplicate of the lectotype: H-NYL!).
The lectotype (L), collected in the Hawaiian Islands, is represented by an apparently burnt
thallus. It contains only atranorin [minor] and chloroatranorin [major] (TLC in solvent C, HPLC
- J.A. Elix, May 2009). The ascospores are ellipsoid, 16-20 x 10-12 um, episporium 1.0-1.5
um. Conidia were not found in this specimen, but the duplicate of the lectotype (H-NYL 35280!)
revealed conidia filiform, 12-16.5 x ca. 1 um. These features exclude this taxon as synonym of
Parmotrema cetratum (Hale & Fletcher 1990), especially the absence of salazinic acid.
Otherwise it can be close to P. macrocarpum (Pers.) Hale (see also Hale & Fletcher 1990).
However, srudies on the type of this species and its synonym Parmelia macrocarpoides Vainio
were not performed yet.
Parmelia menziesii C.W. Dodge
Armais of the Missouri Botanical Garden 46: 101. 1959. - Type: Cape of Good Hope, Menzies
s.n. (holotype: FH-Tuck!). Not Parmelia menziesii Ach., Meth. Lich. 221. 1803 (= Leptogium).
The holotype (FH), from Cape of Good Hope, presents apothecia 1-4 mm in diameter that are
concave or urceolate, stipitate, laminai, with maculate amphithecium, disc brown, epruinose,
imperforate, and immature ascospores ellipsoid 15-18 x 10 um, with episporium ca. 2 um. The
pycnidia are submarginal, conspicuous, without prominent margin, frequent, ostiole black,
conidia filiform, 12-16 x ca. 1.0 um. The córtex is K+ yellow, UV-, and the medulla K+ yellow
— > blood red, C-, KC-, P+ strong yellow, UV-, so attesting the presence of salazinic acid.
The more salient difference of this species is supposedly conidial, but the observations we
made do not match with the protologue (conidia bacillary, 5-6 x 1 um, fide Dodge 1959). The
remaining of the morphology is difficult to evaluate due to the very fragmentary status of the
type, devoid of well developed apothecia, etc. Actually, there are only three parts from one (or
more) thallus glued in a paper and only immature ascospores were found in the larger
apothecium. They revealed similar measures as described by Dodge (1959).
Hale & Fletcher (1990) listed this species as a synonym of Parmotrema cetratum. However,
attention should be placed in the imperforate apothecia, since in Parmotrema cetratum they are
clearly perforate. It is possible they are so because immature, and so it seems convenient to
maintain P. menziesii as a nomen inquirendum until more specimens from the type locality be
studied.
96
Parmotrema moreliense (B. de Lesd.) W.L. Culb. & CF. Culb.
TheBryologist84(3): 310. 1981.
Parmelia moreliensis B. de Lesd., Lichens du Mexique: 5. 1914. - Type: México, Etat de
Michoacan. Morelia; Campanário No 4400 sur écorce (Bouly de Lesdain 1914).
= Everniastrum moreliense (B. de Lesd.) Hale, Mycotaxon 3 (3): 348. 1976.
This type was not located, since the information about the type and where it is located was not
supplied by Bouly de Lesdain (1914), Culberson & Culberson (1981) and Hale (1976).
Parmelia per for ata var. microspora Rãs.
Suom. Eláin.Kasv. Seuran Vanamo 2: 45. 1947. - Type: Argentina, Tucumán, San Xavier,
Herrera 2519 (lectotype: H), fide Hale & Fletcher (1990).
The type could not be found in H. This taxon was listed as synonym of P. cetratam by Hale &
Fletcher (1990).
Lichen perlatus a ciliatus Lam. & DC.
Flore Française 2: 403. 1805- Type: France, Vire, leg. Delise s.n. (lectotype: G, fide Hale &
Fletcher 1990).
= Parmelia perlata g ciliata (Lam. & DC) Duby, Bot. Gall. 2: 601. 1830.
= Parmelia perlata var. tentacalata Wallr., Fl. Crypt. Germ. 3: 521. 1831. Based on Lichen
perlatus a ciliatus.
= Parmelia ciliata (Lam. & DC) Nyl., Flora 61: 247. 1878.
= Parmelia perlata var. ciliata (Lam. & DC) Jatta,F/. Ital. Crypt. 3: 206. 1909.
= Parmelia ciliata (Lam. & DC) Fink, Proc. Indiana Acad. Sei. 1918: 268. 1919. Superíluous
combination, fide Hale & Fletcher (1990).
= Lobaria perlata [beta] ciliata Lam. & DC, fide B. Hale & DePriest (1999).
The lectotype of this species was not found in G. However, the curator (Philippe Clerc) kindly
sent one specimen labeled Parmelia perlata var. ciliata Delise from Vire. This is a well-
developed lichen with ciliate isidia and stictic acid (medulla K+ yellow, P+ orange). The real
identity is beyond the scope of the present work, but it is probably Parmotrema crinitum (A eh.)
M. Choisy.
Since there are no annotations from Hale (or anyone) with the specimen and it clearly do not
belong to Parmotrema reticulatum, the name Lichen perlatus a ciliatus remains to be clarified in
the future.
Parmelia polita Fr.
Syst. Orb. Veg. 1: 283. 1825. - Type: U.S.A., North Carolina, Schwinitz (lectotype: UPS). "As
Mílller Argoviensis (Flora 70: 318. 1887) noted, the published locality (Cape, South Africa)
cited by Fries is incorrect", fide Hale & Fletcher (1990).
The type was not sent by UPS. This taxon was listed as synonym of P. cetratum by Hale &
Fletcher (1990).
97
Excluded name
Parmelinella afrocetrata (Elix, Fischer & Killmann) Marcelli & Spielmann, comb. nov.
Parmotrema afrocetratum Elix, Fischer & Killmann, The Lichenologist 37 (2): 103. 2005. -
Type: Rwanda, Butare, park of the Instituí de Recherche Scientifique et Technologique, c. 1750
m alt., onrocks, 14.X.1999, E. Fischer, s.n. (holotype: B!; isotype: CANB).
(Fig. 82)
Holotype description
Thallus yellowish gray, lobate, loosely adnate, over rocks with debris, 8 cm broad. Lobes
irregularly branched, crowded, 4-8.5 mm wide, surface continuous or irregularly cracked,
smooth, lustrous or opaque, becoming reticulate cracked towards the center; apical zone
rounded; margin smooth to crenate or irregular, undulated. Maculae weak or indistinct, irregular,
laminai. Cilia simple, rare in the axils, very small, up to O.lx 0.02 mm. Lacinules, pustules,
soredia and isidia absent. Medulla white. Under surface black, lustrous, papillate or rugose, with
cracks; marginal zone brown, lustrous, 2-4 mm wide, rhizinate, with an attenuated limit, rugose
or papillate, or rarely with veins; rhizines cone olor to the undersurface, simple or rarely
branched, frequently with penicillate whitish ápices when in the marginal zone, short and thin,
up to 0.5 mm long, 0.01-0.04 mm wide, abundant, evenly distributed.
Apothecia cupuliform, 0.5-2.0 mm in diameter, stipitate, laminai, margin smooth or
sometimes with a profound incision, amphithecium smooth, disc brown, epruinose, imperforate;
ascospores ellipsoid, 8.5-9.0 x 6.0-6.5 um, episporium 0.5-1.0 um. Pycnidia laminai,
conspicuous, usually with prominent margin, abundant, ostiole black; conidia sublageniform,
5.0-7.5 x ca. 1.0 um.
Color tests: córtex K+ yellow, UV-; medulla K+ yellow — > blood red, C-, KC-, P+ strong
yellow, UV-.
TLC in solvent C (J.A. Elix, 21.X.2003, label with the holotype): atranorin (minor), salazinic
acid (major), consalazinic acid (minor).
Remarks: the rarely ciliate (only in the axils) lobes, apothecia imperforate, small ascospores,
conidia sublageniform, medulla with salazinic and consalazinic acid and overall morphology,
including the rhizines that develop a penicillate whitish apex when growing near the marginal
zone, are an indicative that this species do not belong to Parmotrema, being more adequately
treated under Parmelinella Elix & Hale.
Benatti & Marcelli (2007) provided a worldwide key to this genus, and two species with
brown marginal zone could be compared with: P. manipurenses (Singh) Elix & Hale is the only
known saxicolous species, has larger ascospores 9-14 um and smaller conidia 2-A um (Singh
1980), while the corticolous P. simplicior (Hale) Hale & Elix has smaller ascospores 6x4 j^rn
and also shorter conidia 2-3 um (Hale 1972).
This seems the first species in Parmelinella with sublageniform conidia.
Although in the protologue the substrate was described as saxicolous, the holotype has several
wood pieces in the underside, as well mosses and soil. This can indicate the thallus was growing
on debris on rocks.
Distribution: Africa -Rwanda (Elix et ai. 2005, Bock et ai. 2007).
98
Final remarks
The present study of Parmotrema s.l. with salazinic acid resulted in the full type
descriptions and comments on 61 species, two of them new species and 15 new combinations
(01 mParmelinellà).
This shows that the species concept as used nowadays in this group is clearly
unsatisfactory, several good species being recognized as synonyms of few species, said "very
variable" and "widely distributed" in the literature.
Therefore, some important conclusions are:
• maculae, cilia, type and development of soredia and isidia and conidial features constitute,
in Parmotrema with salazinic acid, good specific characters;
• arbuscula were considered structures distinct from isidia;
• distinction was made between reticular maculae, pseudoreticular maculae and
hypermaculate thalli;
• although with recognized importance in the taxonomy of lichenized fungi, conidial features
were usually overlooked in the literature, about 40% of the present conidial information
being described for the first time;
• undersurface features, like brightness, relief and type of rhizines are more important that
previously thought;
• the number of accepted synonyms, especially in tropical species, is very high. However,
when original descriptions are compared, the species seem different; when the types are
compared, there are no doubts that several taxa constitute good species.
The characters and methodology employed seem also applicable to the Parmotrema
without salazinic acid - a good way to test the several statements made here. Only then, some
subgeneric rearrangements or more general conclusions could be established. Anyway,
Parmotrema undoubtedly is a very diverse genus, and a new taxonomical approach is necessary,
especially in the neotropical species.
99
References
ACHARIUS, E. (1803). Methodus qua omnes detectos lichenes. Stockholm. 394 p.
ACHARIUS, E. (1814). Synopsis Methodica Lichenum. Lund. 392 p.
ADLER, M.T. (1988). La família Parmeliaceae (Líquenes, Ascomycotiná) en la Província de
Buenos Aires: Estúdio taxonómico-florístico. Tesis presentada para optar ai título de Doctora
en Ciências Biológicas, Universidad de Buenos Aires. 266 p.
ADLER, M.T. (1992). Clave de los géneros y de las espécies de Parmeliaceae (Lichenes,
Ascomycotiná) de la Provincia de Buenos Aires (Argentina). Boletín de la Sociedad Argentina
de Botânica 28 (1-4): 11-17.
ADLER, M. & J. A. ELIX (1987). Three new saxicolous species in Parmeliaceae (lichenized
Ascomycotiná) from Argentina. Mycotaxon 30: 339-344.
AGUIRRE-C, J. & RANGEL-CH, J.O. (2007). Amenazas a la conservación de las espécies de
musgos y líquenes en Colômbia - una aproximación inicial. Caldasia 29 (2): 235-262.
AHTI, T. & MARCELLI, M.P. (1995). Taxonomy of the Cladonia verticillaris complex in
South America. In: Farkas, E.E., Lílcking, R. & Wirth, V. (eds.): Scripta Licheno lógica -
Lichenological Papers Dedicated to Antonín Vézda. Bibliotheca Lichenologica 58 5-26.
APTROOT, A. (1991). Lichens of Madagáscar: new records and species of Parmeliaceae.
Cryptogamie,Bryol. Lichénol. 12(2): 149-154.
APTROOT, A. (2008). Checklist of lichens and lichenicolous fungi of Madagáscar.
<http: //www.biologie.uni-hamburg . de/checklists/africa/madagascar_l.htm> .
ASAHINA, Y. (1940). Lichenologische Notizen XIV. Journal ofJapanese Botany 16: 592-603.
ASAHINA, Y. (1952). Lichens of Japan. Vol. II. Genus P arme lia. Research Institute for Natural
Resources, Shinjuku, Tokyo. VI + 162 p. + 23 pi.
ASAHINA, Y. & SHIBATA, S. (1954). Chemistry ofLichen Substances. Japanese Society for
the Promotion of Science, Tokyo. VI + 240 p.
AWASTHI, D.D. (1976). Lichen genus Parmelia in índia I - Subgenera Parmelia and
Amphigymnia. Biological Memoirs, Lichenology Series 1: 155-229.
BARBOSA, S.B. (2009). Aplicabilidade taxonômica de variações anatómicas em fungos
liquenizados. Tese apresentada ao Instituto de Biociências, Campus de Botucatu, UNESP,
para obtenção do título de Doutor em Ciências Biológicas (Botânica). 141 p.
BARROS, L.M. de & XAVIER FILHO, L. (1972). Catálogo dos liquens do Herbário do
Departamento de Botânica da Universidade Federal de Pernambuco. Anais do XXIII
Congresso Nacional de Botânica (Garanhuns, PE): 45-55.
BENATTI, M.N. (2005). Os géneros Canomaculina, Parmotrema e Rimelia (Parmeliaceae,
Ascomycetes) no litoral centro-sul do Estado de São Paulo. Dissertação (mestrado). Instituto
de Botânica da Secretaria de Estado do Meio Ambiente. São Paulo. 389 p.
BENATTI, M.N., MARCELLI, M.P. & ELIX, J.A. (2008). Three new species of Parmotrema
containing salazinic acid from the coast of São Paulo State, southeastern Brazil. Mycotaxon
103: 41-52.
BERRY, E.C. (1941). A monograph of the genus Parmelia in North America, North of México.
Annals of the Missouri Botanical Garden 28: 31-146.
100
BLANCO, O., CRESPO, A., DIVAKAR, P.K., ELIX, JA. & LUMBSCH, H.T. (2005).
Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae). Mycologia 97
(1): 150-159.
BOCK, C, HAUCK, M. & FISCHER, E. (2007). The lichen flora of Rwanda: an annotated
checklist. Willdenowia 37: 563-575.
BOULY DE LESDAIN, M. (1914). Lichens du Mexique (états de Puebla et du Michoacan)
recneillis par le frère Arsène Brouard. imp. I. Escalante, S. A., México. 31 pp.
BRODO, LM. (1965) [Review]. M.E. Hale, Jr. A monograph of Parmelia subgenus
Amphigymnia . Contributions from the United States National Herbarium 36 (5): 193-358. 29
fig. 16 pi. The Bryologist 68 (3): 374.
BRODO, LM., SHARNOFF, S.D. & SHARNOFF, S. (2001). Lichens ofNorth America. Yale
University Press, New Haven & London. 795 pp.
BUNGARTZ, F. (2001). Analysis of lichen substances. <http://nhc.asu.edu/lichens/lichen
_info/tlc.jsp> . Accessed in June 2009.
CALVELLO, S. & LIBERATORE, S. (2002). Catálogo de los Líquenes de la Argentina.
Kurtziana 29 (2): 7-170.
CANEZ, L.S. (2005). A família Parmeliaceae na localidade de Fazenda da Estrela, município de
Vacaria, Rio Grande do Sul, Brasil. Dissertação (mestrado). Instituto de Botânica da
Secretaria de Estado do Meio Ambiente. São Paulo. 292 p.
CANÊZ, L.S. & MARCELLI, M.P. (2006). Géneros de Parmeliaceae (Ascomycetes
liquenizados) na localidade de Fazenda da Estrela, Vacaria, Rio Grande do Sul, Brasil.
Caderno de Pesquisa Série Biologia (Santa Cruz do Sul) 18 (3): 38-81.
CENGIA-SAMBO, M. (1930). Licheni delia Patagonia e di altre regioni deli' Argentina raccolti
dai missionari salesiani. Contribua' Scientifici delle Missioni Salesiane dei Beato Don Bosco,
Torino 6: 73 p., 2 maps, 9 pi.
CENGIA-SAMBO, M. (1939a). Lichenes. In: Cufodontis,G. Missione Biológica nel paese dei
Borana. IV. Raccolte Botaniche. R. Acad. Italiana Centro Studi Africa Orient. Ital., p. 375-
389.
CENGIA-SAMBO, M. (1939b). Licheni deli' Africa Orientali Italiana dal Console prof. Senni e
dal Centurione ing. Giordano, Ufficiali delia Milizia Nazionale Forestale di Addis Abeba nel
1937. Nuovo Giornale Botânico Italiano, n. ser. 46 (3): 437-455.
CENGIA-SAMBO, M. (1939c). Licheni dei Brasile. Annali di Botânica 22 (1): 19-41 (1-23).
CHEN, J-B., WANG, S-L. & ELIX, JA. (2003). Parmeliaceae (Ascomycota) lichens in China's
Mainland I. The genera Canomaculina, Parmelina, Parmelinella and Parmelinopsis.
Mycotaxon 86: 19-29.
CHEN, J-B., WANG, S-L. & ELIX, JA. (2005). Parmeliaceae {Ascomycota) lichens in China's
Mainland III. The genusP armo trema. Mycotaxon 91: 93-1 13
CHOISY, M. (1952). Catalogue des lichens de la region lyonnaise. Fase. 9. Bulletin mensuel de
la Société Linnéenne de Lyon et des Société Botanique de Lyon 21: 165-180.
CHRISTENSEN, S.N. & SIPMAN, H.J.M. (1998). Silver moss - ornamental lichens from
Brazil. British Lichen Society Bidletin 82: 11-13.
CULBERSON, W.L. & CULBERSON, CF. (1981). The genera Cetrariastrum and
Concamerella (Parmeliaceae): a chemosystematic synopsis. The Bryologist 84 (3): 273-314.
101
CULBERSON, W.L. (1973). The Parmelia perforata group: niche characteristics of chemical
races, speciation by parallel evolution, and a new taxonomy. The Bryologist 76: 20-29.
DEY, J.P. (1978). Fruticose and foliose lichens of the High-Mountain Áreas of the Southern
Appalachians. The Bryologist 81(1): 1 -93 .
DIVAKAR, P.K. & UPRETI, D.K. (2005). Parmelioid Lichens in índia (a revisionary study).
Bishen Singh Mahendra Pai Singh, Delira Dun, índia. 488 p.
DIVAKAR, P.K., UPRETI, D.K. & ELIX, J.A. (2001). New species and new records in the
lichen family Parmeliaceae (Ascomycotina) from índia. Mycotaxon 80: 355-362.
DIVAKAR, P.K., BLANCO, O. HAWKSWORTH, D.L. & CRESPO, A. (2005). Molecular
phylogenetic studies on the Parmotrema reticulatum (syn. Rimelia reticiãatá) complex,
including the confirmation of P. pseiidoreticiãatam as a distinct species. The Lichenologist 37
(1): 55-65.
DODGE, C.W. (1959). Some lichens of Tropical Africa, III. Parmeliaceae. Annals of the
Missouri Botanical Garden 46: 39-193.
DONHA, CG. (2005). Os géneros Canomaculina, Parmotrema e Rimelia (Ascomycota
liquenizados, Parmeliaceae) na área de proteção ambiental de Guaraqueçaba - Paraná -
Brasil. Dissertação (mestrado). Universidade Federal do Paraná. Curitiba. 118 p.
DUBY, J.E. (1830). Synopsis Plantarum in Flora Gallica descriptarum. 2a ed. Pars II: plantas
cellulares continens. Paris: Desray. p. 542-1068 + 58 p.
DU RIETZ, G.E., (1924a). Die soredien und isidien der Flechten. Svensk Botanisk Tidskrift 18
(3): 371-396.
DU RIETZ, E. (1924b). Kritische Bemerkungen ílber die Parmelia perlata-Gmppe. NytMagazin
for Naturvidenskaberne 62: 63-82.
EGAN, R.S. (1982). Parmotrema arteagum, a new lichen species from México. The Bryologist
85(1): 79-83.
ELIASARO, S. & ADLER, M.T. (1997). Two new species and new reports in the Parmeliaceae
sensu stricto (lichenized Ascomycotina) from Brazil. Mycotaxon 63: 49-55.
ELIASARO, S. & ADLER, M.T. (1998). Rimelia pontagrossensis, a new species in the
Parmeliaceae sensu stricto (Lichenized Ascomycotina) from Brazil. Mycotaxon 66: 127-130.
ELIASARO, S. (2001). Estúdio taxonómico y florístico sobre las Parmeliaceae sensu stricto
(Ascomycota Liquenizados) dei Segundo Planalto dei Estado de Paraná, Brasil. Buenos Aires.
Tesis de Doctor (en Ciências Biológicas). Universidad de Buenos Aires, Facultad de Ciências
Exactas y Naturales. 267 p.
ELIASARO, S. (2006). Checklist of lichens and lichenicolous fungi of Paraná (Brazil). Version
1 September 2006. <http://www.checklists.de>. Accessed in October 2008.
ELIX, J.A. & HALE, M.E. (1987). Canomaculina, Myelochroa, Parmelinella, Parmelinopsis
and Parmotremopsis, five new genera in the Parmeliaceae (Lichenized Ascomycotina).
Mycotaxon 29: 233-244.
ELIX, J.A. (1993). New species in the lichen family Parmeliaceae (Ascomycotina) from
Austrália. Mycotaxon 47: 101-129.
ELIX, J.A. (1994). Parmeliaceae. Flora of Austrália 55: 1-360.
ELIX, J.A. (1997). The lichen genera Canomaculina and Rimeliella (Ascomycotina,
Parmeliaceae). Mycotaxon 65: 475-479.
102
ELIX, J.A. (1998). Clarification of the synonymy and chemistry of Parmotrema zollingeri and
related species. Australasian Lichenology 42: 22-21.
ELIX, J. A. (2001). Additional lichen records from Oceania 7. Parmeliaceae from Fiji.
Australasian Lichenology 48: 34-37.
ELIX, J.A. & JOHNSTON, J. (1988). New species in the lichen family Parmeliaceae
(Ascomycotina) from the Southern Hemisphere. Mycotaxon 31 (2): 491-510.
ELIX, J.A., BAWINGAN, P.A. & FLORES, Y.G. (2002). A new species and further new
records in the lichen family Parmeliaceae {Ascomycotina) from the Philippines. Mycotaxon
81:251-256.
ELIX, J.A., FISCHER, E. & KILLMANN, D. (2005). New saxicolous species of Hypotrachyna
and Parmotrema {Parmeliaceae) from Rwanda. The Lichenologist 37 (2): 101-104.
ESSLINGER, TL. (2008). A cumulative checklist for the lichen- forming, lichenicolous and
allied fungi of the continental United States and Canada. North Dakota State University:
<http://www.ndsu.nodak.edu/instruct/esslinge/chcklst/chcklst7.htm>. (First Posted 1
December 1997, Most Recent Version (#14) 8 October 2008), Fargo, North Dakota.
FEE, A.L.A. (1837). Essai sur les cryptogames des écorces exotiques officinales. 4. Supplément
etRevision. Paris et Strassburg. 180 p + 9 pi.
FERRARO, L.I. (1979). Una nueva espécie de Parmeliaceae para el NE Argentino. Hickenia 1
(34): 191-193.
FERRARO, L.I. & ELIX, J.A. (1993). Two new species of Parmeliaceae (Lichenized
Ascomycota) from South America. Mycotaxon 49: 405-409.
FERRARO, L.I. & ELIX, J.A. (2000). A new species of Canomaculina (Lichenized
Ascomycotina, Parmeliaceae) from Argentina. Mycotaxon 74 (2): 391-394.
FEUERER, T. (1998). E.A. Vainio's contribution to the knowledge of the Parmeliaceae. In:
Marcelli, M.P. & Ahti, T. (eds.), Recollecting Edvard August Vainio. CETESB - Companhia
de Tecnologia de Saneamento Ambiental - Estado de São Paulo, São Paulo, Brazil, p. 47-60.
FEUERER, T. (ed.) (2008). Checklists of lichens and lichenicolous fungi. Version 1 May 2008.
< http: //www.biologie.uni-hamburg . de/checklists/lichens/portalpages/portalpage_checklists_
switch.htm>. Accessed in December 2008.
FEUERER, T., AHTI, T. & VITIKAINEN, O. (1998). Lichenological investigations in Bolivia.
In: Marcelli, M.P. & Seaward, M.R.D. (Eds.). Lichenology in Latin America: history, current
knowledge and applications, p. 71-86. CETESB. São Paulo.
FINK, B. & FUSON, S.C. (1919). Ascomycetes new to the Flora of Indiana. Proceedings ofthe
Indiana Academy of Science: 264-275.
FLEIG, M. (1997). Os géneros Parmotrema, Rimelia e Rimeliella (Lichenes -Ascomycotina,
Parmeliaceae) no Rio Grande do Sul, Brasil. Tese (doutorado em Botânica). Instituto de
Biociências, Universidade de São Paulo. São Paulo, 250 p.
FLEIG, M. & RIQUELME, I. (1991). Liquens de Piraputanga, Mato Grosso do Sul, Brasil. Acta
botânica brasílica 5 (1): 3-12.Fries, E. 1825. Systema Orbis Vegetabilis. Pars I. Plantae
homonemeae. Lund. 374 p.
FRIES, E. (1825). Systema Orbis Vegetabilis. Pars I. Plantae homonemeae. Lund. 374 p.
GALLOWAY, D.J. (1985). Flora ofNew Zealand, Lichens. 662 pp., Wellington, New Zealand.
103
GALLOWAY, D.J. (2007). Flora of New Zealand Lichens. Revised second edition, including
lichen-forming and lichenicolous fungi. Volume II: P aunaria - *Zwackhiomyces. Manaaki
Whenua Press, Lincoln, New Zeland. p. 1007-2261.
GAUDICHAUD-BEAUPRE, M.C. (1826). Voyage autour du monde, execute sur les corvettes
VUranie et la Physcienne, sur les ordres du Cap. L. de Freycinet, en 1817-1820. Botanique.
Paris, 522 p.
GYELNIK, V. (1931). Additamenta ad cognitionem Parmeliarum I. Feddes Repertorium
specierum novarum regni vegetabilis 29: 149-157.
GYELNIK, V. (1934). Lichenes argentinenses a professore C.C. Hosseus collecti. Feddes
Repertorium specierum novarum regni vegetabilis 33: 302-309.
HALE, M.E. (1960). A revision of the South American species of P arme lia determined by
Lynge. Contributions from the United States National Herbarium 36 (1): 1-41.
HALE, M.E. (1965). A monograph of Parmelia subgenus Amphigymnia. Contributions from the
United States National Herbarium 36 (5): 193-358.
HALE, M.E. (1971). Parmelia permaculata, a new lichen from Alabama and México.
Phytologia 21 (6): 425.
HALE, M.E. (1972). Four new species of Parmelia (Lichenes) from índia and the Philippines.
The Bryologist 75 ( 1 ): 97- 1 1 .
HALE, M.E. (1974). New combinations in the lichen genus Parmotrema Massalongo.
Phytologia 28 (4): 334-339.
HALE, M.E. (1976). A monograph of the lichen genus Parmelina Hale (Parmeliaceae).
Smithsonian Contributions to Botany 33: 1-60.
HALE, M.E. (1977). New species in the lichen genus Parmotrema Mass. Mycotaxon 5 (2): 432-
448.
HALE, M.E. (1979). How to know the Lichens. The Pictured-Key Nature Series. Dubuque, Iowa:
Wm. C. Brown Company Publishers. Dubuque. 246 p.
HALE, M.E. (1980). Taxonomy and distribution of the Parmelia flaventior group (Lichens:
Parmeliaceae). Journal ofthe Hattori Botanical Labor atory 47: 75-84.
HALE, M.E. (1983). The Biology of Lichens. Third Edition. Edward Arnold, Baltimore. 190 p.
HALE, M.E. (1990). New species of Parmotrema {Ascomycotina: Parmeliaceae) from Tropical
America. In: Contributions to Lichenology. In honour of A. Henssen. Bibliotheca
Lichenologica 38: 109-1 19.
HALE, M.E. & FLETCHER, A. (1990). Rimelia Hale & Fletcher, a new lichen genus
(Ascomycotina: Parmeliaceae). The Bryologist 93 (1): 23-29.
HALE, B.W. & DePRIEST, P.T. (1999). Mason E. Hale's list of epithets in the parmelioid
genera. The Bryologist 102 (3): 462-544.
HAWKSWORTH, D.L., AHTI, T., VITIKAINEN, O. & CRESPO, A. (2003). (1595) Proposal
to reject the name Parmelia perlata var. olivaria to protect the name Parmotrema
pseudoreticulatum (Ascomycota, Lecanorales, Parmeliaceae). Taxon 52: 626-627.
HERRE, A.W.C.T. (1906). The foliaceous and fruticose lichens of the Santa Cruz Peninsula,
Califórnia. Proceedings ofthe Washington Academy of Sciences 7: 325-396.
HERRE, A.W.C.T. (1910). The lichen flora ofthe Santa Cruz Peninsula, Califórnia. Proceedings
ofthe Washington Academy of Sciences 12 (2): 27-269.
104
HERRE, A.W.C.T. (1942). [Review of] A monograph of the genus Parmelia in North America,
North of México, by Edward C . Berry . The Bryologist 45 (4): 149-151.
HUE, A. (1895). Lichens de Californie récoltés par M. Diguet et determines par M. 1'abbé Hue.
Journal de Botanique 9 (6): 108-113.
HUE, A. (1899). Lichenes extra-europaei a pluribus collectoribus ad Museum Parisiensi missi.
Nouvelles Archives duMuséum d'Histoire Naturelle de Paris, ser. 4(1): 27-220. 6 pi.
HUNECK, S. & YOSHIMURA, I. (1996). Identification oflichen substances. Springer, Berlin.
493 p.
INDEX FUNGORUM (2009). <http://www.indexfungorum.org/Names/Names.asp>.
JATTA, A. (1909-1911). Flora itálica Cryptogama III Lichenes. Tip. CappelliYRocca di S.
Casciano. 958 p.
JUNGBLUTH, P. (2006). A família Pa rmeliaceae (fungos liquenizados) em fragmentos de
cerrados do estado de São Paulo. Dissertação (mestrado). Instituto de Botânica da Secretaria
de Estado do Meio Ambiente. São Paulo. 323 p.
KALB,K. (1982). Lichenes neotropici. Fascikel IV (No. 121-160): 1-12.
KILLMANN, D. & FISCHER, E. (2005). New Records for the Lichen Flora of Rwanda, East Africa.
Willdenowia 35(1): 193-204.
KREMPELHUBER, A. (1869). Parmelia perforata Ach., ihre sichere Erkennung und
Unterscheidung von verwandten Arten. Flora 52: 219-223.
KREMPELHUBER, A. (1873). Lichenes Brasilienses. Partic. XIV von E. Warming, Symbolae
ad floram Brasiliae cognoscendum. Videnskabelige meddelelser fra den Naturhistoriske
forening i Kjòbenhavn 5: 1-35 + 1 plate.
KREMPELHUBER, A. (1876a). Lichenes brasilienses collecti a D. A. Glaziou in provincia
brasiliensi Rio Janeiro. Flora (Regensburg) 59: 33-38, 56-63, 70-77, 140-144, 172-176, 217-
224, 248-251, 265-271, 316-320, 378-384, 412-416, 417-423, 445-482, 475-482, 511-514,
522-529.
KREMPELHUBER, A. (1876b). Neue Beitráge zur Flechtenflora Neu-Seelands. Verhandlungen
der zoologisch-botanischen Gesellschaft in Wien 26: 447-460.
KREMPELHUBER, A. (1877). Neue Beitráge zu Afrika's Flechten-flora. Linnaea 7: 135-144.
KREMPELHUBER, A. (1878). Lichenes, collecti in republica Argentina a Doctoribus Lorentz et
Hieronymus. Flora (Regensburg) 61: 433-439, 461-464, 476-480, 492-496, 516-523.
KREMPELHUBER, A. (1881). Ein neuer Beitrag zur Flechten-Flora Australiens.
Verhandlungen der zoologisch-botanischen Gesellschaft in Wien 30: 329-342.
KROG, H. (1974). Parmelia ultralucens, a new lichen species in subgenus Amphigymnia. The
Bryologist 11 (2): 253-256.
KROG, H. & SWINSCOW, T.D.V. (1981). Parmelia subgenus Amphigymnia in East Africa.
Bulletin of the British Museum (Natural History) Botany series 9 (3): 143-231.
KROG, H. & SWINSCOW, T.D.V. (1983). A new species and new combinations in
Parmotrema (Parmeliaceae). The Lichenologist 15 (2): 127-130.
KUROKAWA, S. (1974). Four new species of Parmelia from Brazil. Bulletin of the National
Science Museum Tokyo, 17 (4): 297-300 + 1 pi.
105
KUROKAWA, S. (1979). Enumeration of species of Parmelia in Papua New Guinea. In: S.
Kurokawa (ed.): Studies on Cryptogams of Papua New Guinea. Academic Scientific Book
Inc., Tokyo, pp. 125-148.
KUROKAWA, S. (1991). Japanese species and genera of the Parmeliaceae. Journal ofJapanese
Botany 66 (3): 152-159.
KUROKAWA, S. (2001). Taxonomic notes on Parmelia reparata (Parmeliaceae, Lichenes) and
the related species. Bulletin ofthe National Science Museum Tokyo, Ser. B, 27 (1): 1-10.
KUROKAWA, S. (2003). Checklist of Japanese Lichens. National Science Museum, Tokyo.
128 p.
KUROKAWA, S. & LAI, M-J. (2001). Parmelioid lichen genera and species in Taiwan.
Mycotaxon 77: 225-284.
LAMARCK, M.M. de & de CANDOLLE, A.P. (1805). Flore Francaise. Troisième edition. Vol.
II. Paris. 600 p.
LAMB, LM. (1963). Index nominum lichenum. Inter annos 1932 et 1960 divulgatorum. Ronald
Press Company, New York. 809 p.
LENDEMER, J.C. & HEWITT, D. A. (2002). A catalogue ofthe type specimens of lichens in the
Herbarium ofthe Academy of Natural Sciences of Philadelphia. Proceedings of the Academy
of Natural Sciences ofPhyladelph ia 1 52 : 1 73 -204 .
LINDSAY, L.W. (1866). Observations on New Zealand Lichens. Transactions of the Linnean
Society ofLondon 25: 493-560.
LOUWHOFF, S.H.J.J. & ELIX, J.A. (1998). The lichen family Parmeliaceae (Ascomycotina) on
Lord Howe Island, Austrália. Mycotaxon 68: 429-463.
LOUWHOFF, S.H.J.J. & ELIX, J.A. (1999). Parmotrema and allied lichen genera in Papua New
Guinea. Bibliotheca Lichenologica 73: 1-152.
LOUWHOFF, S.H.J.J. & ELIX, J.A. (2000). Five new species in the lichen family Parmeliaceae
(Ascomycotina) from Grande Terre, New Caledónia. Mycotaxon 75: 195-203.
LYNGE, B. (1914). Die Flechten der ersten Regnellschen Expedition. Die gattungen
Pseudoparmelia gen. nov. und Parmelia Ach. Arkiv for Botanik 13 (13): 1-172.
LYNGE, B. (1924). On some South American lichens of the Genera Parmelia, Candelária,
Theloschistes and Pyxine. Nyt Magazin for Naturvidenskaberne 62: 83-97.
MACKAY, J.T. (1836). Flora Hibernica, comprising the Flowering Plants, Characeae, Musci,
Hepaticae, Lichenes andAlgae oflreland. Dublin, London & Edinburgh. 650 p.
MAGNUSSON, A.H. & ZAHLBRUCKNER, A. (1944). Hawaiian Lichens. II. The Families
Lecideaceae to Parmeliaceae. Arkiv for Botanik 31A (6): 1-109.
MALCOLM, W.M. & D.J. GALLOWAY, (1997). New Zealand Lichens. Checklist, Key, and
Glossary. Museum of New Zealand Te Papa Tongarewa, Wellington, 192 p.
MARCELLI, M.P. (1991). Aspects ofthe foliose lichen flora ofthe southern-central coast of São
Paulo State, Brazil. In: Galloway, D.J. (ed.). Tropical Lichens: Their Systematics,
Conservation, and Ecology. Systematics Association Special Volume n° 43: 151-170.
Clarendon Press. Oxford.
MARCELLI, M.P. (1992). Ecologia liquênica nos Manguezais do Sul-Sudeste Brasileiro.
Bibliotheca lichenologica 47: 1-VII + 288 p. + 31 photos in 22 pages.
106
MARCELLI, M.P. (2008). Checklist of lichens and lichenicolous fungi of Brazil. Preliminary
version 1 May 2008. <http://www.biologie.uni-hamburg.de/checklists/lichens/south-america/
brazil_l.htm>.
MARCELLI, M. P. & T. AHTI (eds.) (1998). Recollecting Edvard August Vainio. São Paulo:
CETESB. 188p.
MARCELLI, M.P. & BENATTI, M.N. (2008). Espécies de Parmotrema {Parmeliaceae,
Ascomycetes liquenizados) com rizinas dimórficas do litoral centro-sul do Estado de São
¥&u\o.Hoehnea35>{2): 171-183.
MARCELLI, M.P. & RIBEIRO, C.H. (2002). Twenty-one new species of Parmeliaceae
(lichenized fungi) from southeastern Brazil. Mitteihmgen aus dem Instituí fiir Allgemeine
BotanikHamburg 30-32: 125-155.
MARCELLI, M.P., JUNGBLUTH, P., BENATTI, M.N., SPIELMANN, A.A., CANÊZ, L.S.,
CUNHA, I.P.R. & MARTINS, M.F.N. (2007). Some new species and combinations of
Brazilian lichenized fungi. Bibliotheca Lichenologica 96: 209-227.
MASSALONGO, A (1860). Esame comparativo di alcuni generi di licheni. Atti delVIstituto
Veneto di scienze, lettere ed arti 5 (3): 247-267, 313-337.
MATA GARCIA, L.B. (1994). Rimelia bonplandii, a new lichen species (Ascomycotina:
Parmeliaceae). The Bryologist 97 (4): 432-433.
McNEILL, J., BARRIE, F.R., BURDET, H.M., DEMOULIN, V., HAWKSWORTH, D.L.,
MARHOLD, K., NICOLSON, D.H., PRADO, J., SILVA, P.C., SKOG, J.E., WIERSEMA,
J.H. & TURLAND, N.J. (editors & compilers) (2007). International Code of Botanical
Nomenclature (Vienna Code) adopted by the Seventeenth International Botanical Congress
Vienna, Áustria, July 2005.Gantner, Ruggell. (Regnum Vegetabile 146). XVIII, 568 p.
MERRILL, G.K. (1909). Lichen notes, no. 9: Parmelia latíssima Fée and two commonly
associated species. The Bryologist 12: 29-31.
MEYEN, J. & FLOTOW, J. (1843). Lichenes . Nova Acta Acad. Caesar. Leopold.-Carol 19,
Suppl.: 209-232.
MOON, K.H., KUROKAWA, S. & KASHIWADANI, H. (2001). The genus Rimelia (Lichens)
from the Hawaiian Islands. Journal ofJapanese Botany 76 (6): 321-328.
MOORE, B.J. (1968). The Macrolichen Flora of Florida. The Bryologist 71 (3): 161-267.
MORTON, J.B., BENTIVENGA, S.P & WHEELER, W.W. (1993). Germ plasm in the
International Collection of Arbuscular and Vesicular-arbuscular Mycorrhizal Fungi (INVAM)
and procedures for culture development, documentation and storage. Mycotaxon 48: 491-528.
MÚLLER ARGOVIENSIS, J. (1880). Lichenes Africae occidentalis a cll. Dr. Pechuel-Loesche
et Soyaux e regione íluminis Quillu et ex Angola missi, in Mus. bot. reg. berolinensi servati.
Linnaea 9: 31-48.
MÚLLER ARGOVIENSIS, J. (1883). Lichenologische Beitráge 17. Flora (Regensburg) 66: 17-
25,45-48,75-80.
MÚLLER ARGOVIENSIS, J. (1886). Lichenologische Beitráge 23. Flora (Regensburg) 69:
124-128.
MÚLLER ARGOVIENSIS, J. (1888). Lichenologische Beitráge 27. Flora {Regensburg) 71: 17-
25,44-48.
MÚLLER ARGOVIENSIS, J. (1891a). Lichenologische Beitráge 35. Flora (Regensburg) 74:
371-382.
107
MÚLLER ARGOVIENSIS, J. (1891b). Lichenes Schenckiane a cl. Dr. H. Schenck, Bonnensi, in
Brasiliae orientalis prov. Santa Catharina, Paraná, Rio de Janeiro, Minas Geraes et
Pernambuco lecti. Hedwigia 30: 219-234.
MÚLLER ARGOVIENSIS, J. (1892). Lichenes exotici Herbarii Vindobonensis - 1. Lichenes in
Austrália et in ejus vicinitate lecti. Annalen des K. K. naturhistorischen Hofmuseums 7: 302-
305.
MÚLLER ARGOVIENSIS, J. (1893). Lichenes, in Durand, Th. & Pittier, H., Primitiae florae
Costaricensis. Séconde énumération. Bulletin de la Société royale de Botaniqae de Belgique
32: 122-173.
MÚLLER ARGOVIENSIS, J. (1894). Lichenes usambarenses. Engler's Botanische Jahrbilcher
20: 238-298.
NASH III, T.H. & ELIX, J.A. (2002a). Canomaculina. In: Nash III, T.H., Ryan, B.D., Gries, C.
& Bungartz, F. (eds.). Lichen Flora of the greater Sonoran Desert Region. Volume 1. Lichens
Unlimited, Arizona State University, Tempe, Arizona, USA. p. 120-122.
NASH III, T.H. & ELIX, J.A. (2002b). Parmotrema. In: Nash III, T.H., Ryan, B.D., Gries, C. &
Bungartz, F. (eds.). Lichen Flora of the greater Sonoran Desert Region. Volume 1. Lichens
Unlimited, Arizona State University, Tempe, Arizona, USA. p. 318-329.
NASH III, T.H. & ELIX, J.A. (2002c). Rimelia. In: Nash III, T.H., Ryan, B.D., Gries, C. &
Bungartz, F. (eds.). Lichen Flora of the greater Sonoran Desert Region. Volume 1. Lichens
Unlimited, Arizona State University, Tempe, Arizona, USA. p. 449-451 .
NYLANDER, W. (1869). Lichenes in Brasilia a Glaziou collecti. Flora (Regensburg) 52: 117-
126.
NYLANDER, W. (1875). Liste des lichens recueilles par M.G. de 1'Isle, aux iles Saint-Paul et
d'Amsterdam et description des espèces nouvelles. Compt Rend. Hebd. Séances Académie
des Sciences [Paris] 81: 725-726.
NYLANDER, W. (1878). Addenda nova ad Lichenographiam Europaeam. Continuatio
tricesima. Flora (Regensburg) 61: 241-249.
NYLANDER, W. (1885). Parmeliae exoticae novae. Flora (Regensburg) 68: 605-615.
NYLANDER, W. (1886). Lichenes Insulae Sancti Pauli. Flora (Regensburg) 69: 318-322.
NYLANDER, W. (1900). Lichenes Ceylonenses et Additamentum ad Lichenes Japoniae. Acta
Societatis Scientiarum Fennicae 26 (10): 1-33.
ORANGE, A., JAMES, P.W. & WHITE, F.J. (2001). Microchemical methods for the
identification of lichens. British Lichen Society. 101 p.
OSÓRIO, H.S. (1970). Lichens from Cantera, South Paraguay. Comunicaciones Botânicas dei
Museo de Historia Natural de Montevideo 4 (50): 1-3.
OSÓRIO, H.S. (1972). Contributions to the Lichen Flora of Uruguay. VIL A preliminary
catalogue. Comunicaciones Botânicas dei Museo de Historia Natural de Montevideo 4 (56):
1-46.
OSÓRIO, H.S. (1976). Contribution to the Lichen Flora of Argentina VIII. Lichens from Punta
Lara, Buenos Aires Province. The Bryologist 79 (3): 358-360.
OSÓRIO, H.S. (1977a). Contribution to the Lichen Flora of Brazil. II. Lichens from
Guarapuava, Paraná State. Dusenia 10 (2): 101-102.
OSÓRIO, H.S. (1977b). Contribution to the Lichen Flora of Brazil III. Lichens from Western
Paraná. Acta Biológica Paranaense 6 (1-4): 3-7.
108
OSÓRIO, H.S. (1979). Contribution to the lichen flora of Uruguay XI. Lichens from Arazati,
San José Department. The Bryologist 82 (3): 472-473.
OSÓRIO, H.S. (1981). Contributions to the lichen flora of Argentina XIII. Lichens from
Misiones Province. Communicaciones Botânicas dei Museo de Historia Natural de
Montevideo 4 (63): 1-18.
OSÓRIO, H.S. (1992a). Contribution to the Lichen Flora of Brazil. XXIX. Lichens from Ponta
Porá, Mato Grosso do Sul. Comunicaciones Botânicas dei Museo de Historia Natural de
Montevideo 5 (98): 1-6.
OSÓRIO, H.S. (1992b). Contribución a la flora liquénica dei Uruguay. XXV. Líquenes
publicados entre 1 972 a 1991 . Anales dei Museo Nacional de Historia Natural de Montevideo
2a Serie, Vol. 8: 43-70.
OSÓRIO, H.S. (1995). Contribution to the Lichen Flora o f Uruguay. XXVIII. Lichens from
Southern Rocha. Comunicaciones Botânicas dei Museo de Historia Natural de Montevideo 5
(103): 1-12.
OSÓRIO, H.S. (1997). Contribution to the Lichen Flora of Brazil. XXXIV. Lichens from
Lagura, Santa Catarina State. Comunicaciones Botânicas dei Museo de Historia Natural de
Montevideo 6 (108): 1-4.
OSÓRIO, H.S. (2003). Contribution to the lichen flora of Uruguay. XXXVIII. Some collections
from Eastern Uruguay. Comunicaciones Botânicas dei Museo de Historia Natural de
Montevideo 6 (126): 1-11.
PARK, Y.S. (1990). The Macrolichen Flora of South Korea. The Bryologist 93 (2): 105-160.
PURVIS, O.W., COPPINS, B.J., HAWKSWORTH, D.L., JAMES, P.W. & MOORE, D.M.
(Eds.) (1992). The Lichen Flora of Great Britain and Ireland. London: Natural History
Museum. 710 p.
RÀSÀNEN, V. (1938). Beitráge zur Flechtenflora Sudamerikas. Uruguay ische Flechten,
gesammelt von W.G. Herter. Mit Berílcksichtigung von Material aus anderen Lãndern.
Revista Sudamericana de Botânica 5 (3-4): 65-72.
RÀSÀNEN, V. (1944). Lichenes novi I. Annales Botanici Societatis Zoologicae Botanicae
Fennicae Vanamo 20 (3): 1-34.
RÀSÀNEN, V. (1947). Lichenes novi III. Archivum societatis zoologicae botanicae fennicae
Vanamo 2: 45-51.
RIBEIRO, C.H. (1998). A família Parmeliaceae {Ascomycota liquenizados) em regiões
montanhosas dos estados de Minas Gerais, Rio de Janeiro e São Paulo. Dissertação (mestrado
em Botânica). Instituto de Biociências, Universidade de São Paulo. 194 p.
SINGH, K.P. (1980). A new species oíParmelina from índia. The Bryologist 83 (4): 533-534.
SIPMAN, H.J.M. (1999). Checklist of the lichens of the Guianas.
<http://botany.si.edu/BDG/aboutlich.html>
SIPMAN, H.J.M. (2005). Mason Hale's key to Parmotrema, revised edition: key to wide-lobed
parmelioid species occurring in Tropical America (genera Canomaculina, Parmotrema,
Rimelia, Rimeliellá). Botanischer Garten und Botanisches Museum Berlin-Dahlem, Freie
Universitát Berlin. <http://www.bgbm.org/sipman/keys/Neoparmo.htm>. Accessed in April
2009.
SIPMAN, H.J.M. & van AUBEL, R.J.M.T. (1992). New Parmeliaceae (Lichenes) from the
Guianas and surroundings. Mycotaxon 44 (1): 1-12.
109
SMITH, C.W. (1993). Notes on Hawaiian Parmelioid Lichens. The Bryologist 96 (3): 326-332.
SPIELMANN, A.A. (2005). A família Parmeliaceae (fungos liquenizados) nos barrancos e
peraus da encosta da Serra Geral, Vale do Rio Pardo, Rio Grande do Sul, Brasil. Dissertação
(mestrado). Instituto de Botânica da Secretaria de Estado do Meio Ambiente. São Paulo. 204
P-
SPIELMANN, A.A. (2006). Checklist of lichens and lichenicolous fungi of Rio Grande do Sul
(Brazil). Caderno de Pesquisa Série Biologia (Santa Cruz do Sul) 18 (2): 7-125.
SPIELMANN, A.A. & MARCELLI, M.P. (2006). Preliminary world-wide key to the lichen
genus Rimelia {Parmeliaceae, lichenized Ascomycotá) In: Anais da Primeira Reunião
Brasileira de Estudos Liquenológicos (la REBEL). São Paulo - SP: Instituto de Botânica
(Grupo de Estudos Liquenológicos, GEL-IBt), v.l: 59-69.
SPIELMANN, A.A. & MARCELLI, M.P. (2009). Parmotrema s.l. {Parmeliaceae, lichenized
Ascomycotá) from Serra Geral slopes in central Rio Grande do Sul State, Brazil. Hoehnea.
Accepted for publication.
STAFLEU, F.A., BONNER, C.E.B, McVAUGH, R., MEIKLE, R.D., ROLLINS, R.C., ROSS,
R., SCHOPF, J.M., SCHULZE, G.M., DE VILMORIN, R. AND VOSS, E.G. (Eds.) (1972).
International Code of Botanical Nomenclature, adopted by the Eleventh International
Botanical Congress, Seattle, August 1969.
STIRTON, J. (1877). Additions to the lichen flora of South Africa. Transactions ofthe Glasgow
Society ofField Naturalists 5:21 1-220.
STIRTON, J. (1877-1878). On certain lichens belonging to the genus Parmelia. Scottish
Naturalist 4: 200-203, 252-254, 298-299.
STREIMANN, H. (1986). Catalogue of the Lichens of Papua New Guinea and Irian Jaya.
Bibliotheca Lichenologica 22: 1-145.
SWINSCOW, T.D.V. & KROG, H. (1988). Macrolichens of East Africa. British Museum
(Natural History), London. 390 p.
TAVARES, C.N. (1945). Contribuição para o estudo das Parmeliáceas Portuguesas. Portugaliae
Acta Biológica, série B. 1(1-2): 1-210.
TAYLOR, T. (1847). New lichens, principally from the Herbarium of Sir William J. Hooker.
The London Journal ofBotany 6: 148-197.
TAYLOR, C.J. (1967). The lichens of Ohio. Part I Foliose Lichens. The Ohio Biological
Survey. Biological Notes No. 3. 15 lp.
TENÓRIO, L.U., SIPMAN, H.J.M. & LUCKING, R. (2002). Preliminary checklist of lichens
from Costa Rica. Version 1.2 (June 2002). <http://www.fieldmuseum.org/ticolichen/
checklist.html>. Accessed in April 2009.
VAINIO, E.A. (1890). Étude sur la classification naturelle et la morphologie des Lichens du
Brésil. Pars prima. Acta Societatis pro Fauna et Flora Fennica 7 (1): I-XXIX, 1-247.
VAINIO, E.A. (1909a). Lichenes. In: Schmidt, J. Flora of Koh Chang. Contributions to the
knowledge ofthe vegetation ofthe Gulf of Siam. Part. IX. Botanisk Tidsskrift 29: 104-152.
VAINIO, E.A. (1909b). Lichenes insularum Philippinarum, I. Philippine Journal of Science C,
Bot, IV, 5: 651-662.
VAINIO, E.A. (1918). Lichenes ab A. Yasuda in Japonia collecti. Botanical Magazine [Tokyo]
32: 154-163.
110
VARESCHI, V. (1962). Resultados liquenológicos de excursiones efectuadas en Venezuela 2.
Acerca dei género Parmelia en Vemezuela. Acta Biológica Venezuelica 3 (14): 201-232.
VARESCHI, V. (1973). Resultados licheno lógicos de excursiones effectuadas en Venezuela. No.
3. Catalogo de los liquenes de Venezuela. Acta Botânica Venezuelica 8 (1-4): 177-245.
WALLROTH, F.W. (1831). Flora cryptogamica Germaniae . Norimbergae. 654 p.
WHITE, F.J. & JAMES, P.W. (1985). A new guide to microchemical tecniques for the
identification of the lichen substances. Britsh Lichen Society Bulletin 57 (suppl.): 1-41.
WIGGINS, I.L. (1962). Albert William Christian Theodore Herre (1868-1962). The Bryologist
65 (4): 268-277.
WINNEM, B. (1975). Parmelia subgenus Amphigymnia in Ethiopia. Norwegian Journal of
Botany 22: 139-166.
ZAHLBRUCKNER, A. (1902). Studien íiber brasilianische Flechten. Sitzungsberichte der
kaiserlichen Akademie der Wissenschaften Wien, Mathematisch-Naturwissenschaftlichen
Classe 111, Abtheilung 1: 357-432 + tafel I-II.
ZAHLBRUCKNER, A. (1904). Lichenes a cl. Damazio in montibus Serra do Ouro Preto
Brasiliae lecti in herb. Barbey-Boissier asservati. Bulletin de L 'Herbier Boissier Ser. 2, 4:
134-136.
ZAHLBRUCKNER, A. (1905). Lichenes, a cl. Damazio in Brasilia lecti. II. Bulletin de
L 'Herbier Boissier Ser. 2, 5: 539-543.
ZAHLBRUCKNER, A. (1907). Lichenes (Flechten). Spezieller Teil, p. 49-249. In: Engler, A. &
Prantl, K. (eds.). Die natiirlichen Pflanzenfamilien, sect. 1.1. Leipzig.
ZAHLBRUCKNER, A. (1908). Neue Flechten, IV. Annales Mycologici 6: 129-134.
ZAHLBRUCKNER, A. (1909). Lichenes. In: Schiffner, V. (ed.). Ergebnisse der botanischen
Expedition der Kaiserlichen Akademie der Wissenschaften nach Síidbrasilien 1901. Band II:
Tallophyta und Bryophyta. Denskschriften der Mathematsch-Naturwissenschaftlichen Klasse
der Kaiserlichen Academie der Wissenschaften 83: 87-21 1.
ZAHLBRUCKNER, A. (1926). Afrikanische Flechten (Lichenes). Botanische Jahrbucher fur
Systematik, Pflanzengeschichte und Pflanzengeographie 60: 468-552.
ZAHLBRUCKNER, A. (1928). Neue und ungenílgend beschriebene javanische Flechten.
Annales de Cryptogamie Exotique 1: 109-212.
ZAHLBRUCKNER, A. (1930a). Catalogus lichenum universalis. Band VI. Leipzig: Gebrílder
Borntraeger. 618 p.
ZAHLBRUCKNER, A. (1930b). Lichenes (Ubersicht íiber sámtliche bisher aus China bekannten
Flechten). In: Handel-Mazetti,H: Symbolae sinicae. Botanische Ergebnisse der Expedition
der Akademie der Wissenschaften in Wien nach Siidwest-China 1914-1918. III. J. Springer,
Wien. 254 p. + 1 pi.
ZAHLBRUCKNER, A. (1934). Catalogus lichenum universalis. Band IX. Gebrílder
Borntraeger, Leipzig. 606 p.
111
Figures 1-2: 1 - Holotype of Parmotrema acanthifolium (P); 2 - Part of the holotype of
P. albinatam (TNS). Scales in millimeters.
112
Figures 3-4: 3 - Part of the holotype oiParmotrema albinatum (TNS), underside; 4 - Holotype
oíP. aastrocetratiim (CHR). Scales inmillimeters.
113
Figures 5-6: 5 - Holotype oíParmotrema aiistrocetratum (CHR), upper surface with schizidia;
6 - Holotype of P. austromaciilatam (SP). Scales in millimeters.
114
Figures 7-9: 7 - Holotype of Parmotrema bifidum (S); 8 - Holotype of P. bifidum,
undersurface; 9 - Holotype of P. bifidum, furcated cilia. Scales in millimeters, except as
indicated.
115
li ii i li i í 1 1 li 1 1 1 1 1 ii i iií ;i 1 1 ih 1 1 ii i
Figures 10-11: 10 - Holotype oíParmotrema bifidum (S), upper surface with strong effigurate
maculae; 11 - Lectotype of P. cetratam (H-ACH). Scales in millimeters, except as indicated.
116
Figures 12-13: 12 - Part of the lectotype of P armo trema clavuliferiim (H); 13 - Lectotype of
P. concors (M). Scales in millimeters.
117
Figures 14-15: 14 - Part of the holotype of Parmotrema concurrens (US); 15 - Part of the
holotype of P. coralliforme (US), showing arbuscular structures (arrows). Scales in millimeters.
118
£§Ílb tê
Figures 16-17: 16 - Lectotype of P armo trema cristifenim (FH); 17 - Holotype of P arme lia
mesotropa f. sorediosa (G). Scales in millimeters.
119
I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I ! I I I I I I I I I I I
Figures 18-19: 18 - Holotype of Parmotrema cristobaliae (CTES); 19 - Holotype of
P. delicatulum (TUR-V). Scales in millimeters.
120
I I I I I I I I I I I I I I I I I I I I I ! I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I
Figures 20-21: 20 - Part of the isotype of Parmotrema elixii (CANB); 21 - Isotype of
P. enteroxanthum (UPS). Scales in millimeters.
121
Figure 22: Lectotype oíParmotrema embescens (BM). Scale as indicated.
122
Parmelia eurysaca Htte HOLOTTES
Atranorine, salacinic acià +
(Meà. K+ rub. p+)
det. Mason E. Hale, Jr., U. S. National Museum 1 QÇQ
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Figure 23: Holotype ofParmotrema eurysacum (P). Scale in millimeters.
123
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. ■ • «
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Figures 24-25: 24 - Part of the holotype of Parmotrema expansum (US); 25 - Holotype of
P. flavescens (M). Scales in millimeters.
124
Figures 26-27: 26 - Holotype of Parmotrema foliolosum (FH); 27 - Holotype of P. foliolosum,
showing the undersurface. S cales in millimeters.
125
- ff.-
Figure 28: Holotype of P armo trema granulare (TNS). Scale in millimeters.
28
126
Figures 29-30: 29 - Holotype oíParmotrema herrei (FH); 30 - Holotype of P. lacteum (SP).
S cales in millimeters.
127
Espèces:
í/rrri/es ce?nime rt<?uve//es*. f.í ,
Figures 31-32: 31 - Lectotype of Parmotrema latissimiim (G); 32 - Holotype of
P. leucosemothetum (P). Scales in millimeters.
128
•
1^1%
P^^_
\
■ ^«^.
.'■■■
f
*
®
1 mm
^ i
Figures 33-34: 33 - Holotype of P leucosemothetiim, upper surface with strong effigurate
maculae; 34 - Holotype of P. leucosemoihetum, under marginal zone. Scales indicated.
129
Figures 35-36: 35 - Holotype of Parmotrema lichexanthoniciim (UPCB); 36 - Lectotype of
P. livido-tesselatiim (P). Scales in millimeters.
130
Figures 37-38: 37 - Holotype of Parmotr ema magmim (S); 38 - Isotype of P. magnum (LD),
showing the rugose and lacerate apothecia. Scales in millimeters or as indicated.
131
Figures 39-40: 39 - Holotype of Parmelia microdactyla (UPS); 40 - Holotype of
P. microdactyla, showing the lobules (arrows). Scales in millimeters.
132
S^'í'%
Figures 41-42: 41 - Isotype oíParmotrema mantiqueirense (US); 42 - Part of the lectotype of
P. margaritatum (P). Scales in millimeters.
133
Figures 43-44: 43 - Holotype of Parmotrema masonii (CTES); 44 - Part of the holotype of
P. maximum (P). Scales in millimeters.
134
.» ■,
JS^-Ms -
"'*\
>. **\
Figures 45-46: 45 - Part of the holotype of P armo trema mirandum (US); 46 - Holotype of
P. neotropiciim (US). Scales in millimeters.
135
Figures 47-48: 47 - Part of the holotype of Parmotrema nudum (P); 48 - Holotype of
P. nylanderi (S). Scales in millimeters.
136
^,\/*-
.**
1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
Figures 49-50: 49 - Holotype of Parmotrema parahypotropnm (TNS); 50 - Holotype of
P. parahypotropum, showing the soralia. Scales in millimeters, except as indicated.
137
Figures 51-52: 51 - Holotype of Parmotrema paramoreliense (DUKE); 52 - Holotype of
P. paramoreliense , showing effigurate maculae and the black marginal line. Scales in
millimeters, except as indicated.
138
I I
Figures 53-54: 53 - Part of the holotype of Parmotrema pectinatum (SP); 54 - Holotype of
P. permaciilatam (US). Scales in millimeters.
139
Figures 55-56: 55 - Part of the holotype oí Parmotrema petropoliense (W); 56 - Holotype of
P. petropoliense, showing cilia and soralia. Scales in millimeters, except as indicated.
140
Figures 57-58: 57 - Holotype of Parmotrema pontagrossense (UPCB); 58 - Syntype of
P. pseudoreticulatum (F). Scales in millimeters.
141
s
***;4P
Figures 59-60: 59 - Lectotype of Parmotrema radiatam (S); 60 - Holotype of P. ramusculum
(BM). Scales in millimeters.
142
Figures 61-62: 61 - Holotype of Parmotrema reitzii (US); 62 - Holotype of P. reitzii, showin£
the undersurface. Sc ales in millimeters.
143
Figures 63-64: 63 - Lectotype of Parmotrema reparatum (BM); 64 - Holotype of
P. reterimiãosiim (WU). Scales in millimeters.
144
Figures 65-66: 65 - Duplicate of the lectotype of Parmotrema reticulatum (BM); 66 -
Lectotype of P. ruminatum (W). Scales in millimeters.
■<&>■
145
li
mi^ m
*V A
>
y
(67,
MM
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* -iÓ*)* &
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Figures 67-68: 67 - Holotype of Parmotrema riiptum (S); 68 - Part of the holotype of
P. sieberi (FH). Scales in millimeters.
146
Figures 69-70: 69 - Holotype of Parmotrema spinibarbe (TNS); 70 - Holotype of
P. spinibarbe, showing the cespitose cilia. S cales in millimeters, except as indicated.
147
I I I II I I I I II I I I I I I I
Figures 71-72: 71 - Isotype of Parmotrema spinibarbe (G), showing the botryose structure;
72 - Holotype of P. stuppeum (FH). Scales in millimeters, except as indicated.
148
^_
Figures 73-74: 73 - Holotype of Parmotrema subcaperatum (M); 74 - Lectotype of
P. subisidiosnm (G). Scales in millimeters.
149
Figures 75-76: 75 - Holotype of Parmotrema subsumptum (H-NYL); 76 - Holotype of
P. subtinctorhim (WU). S cales in millimeters.
150
^í-'-
Figures 77-79: 77 - Holotype of Parmotrema ultralucens (O); 78 - Holotype of
P. verrucisetosum (B). 79 - Holotype of P. verrucisetosiim, showing ciliate warts. Scales in
millimeters, except as indicated.
151
1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
Figure 80: Lectotype oíParmotrema warmingii (M). Scales in millimeters.
152
Figures 81-82: 81 - Lectotype of P arme lia badia (P); 82 - Holotype of Parmelinella
afroce trata (B). Scales in millimeters.
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